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Alexander Schintlmeister

This book is the first comprehensive work on oriental Notodontidae (Lepidoptera) outside mainland Asia. The studied area includes also Borneo Island, the Malayan Peninsula, entire New Guinea with adjacent islands. All species are illustrated in both sexes with a total number of 1272 specimens on 51 colour plates. Genitalia photos of both sexes as well as detailed distribution maps are provided for each species.
The book deals in the first volume with 298 species and contains descriptions of 99 new notodontid taxa. A second volume will treat with the remaining 160 species and include also a comprehensive biogeographic analysis.

Atlas of the Hoverflies of Greece

(Diptera: Syrphidae)

Ante Vujić, Martin Speight, Michael Edwin de Courcy Williams, Santos Rojo, Gunilla Ståhls, Snežana Radenković, Laura Likov, Marija Miličić, Celeste Pérez-Bañón, Steven Falk and Theodora Petanidou

The Atlas of the Hoverflies of Greece is the first of a kind within the Mediterranean region. It is the result of decades of research, many travels into the fascinating habitats of Greece (a biodiversity hotspot), visits to world museums, and many people’s passion for hoverflies.
The Atlas is a concise presentation of all 418 hoverfly species for Greece known so far. The species are documented with photos and distribution GIS-maps and they are preceded by a general introduction on the hoverflies and Greek nature, and a generic key.
The Atlas of the Hoverflies of Greece is a handbook for insect aficionados, students and teachers, everyone interested in nature, and managers and conservationists aiming at raising public awareness of a nature nowadays threatened more than ever.


Judith Benz-Schwarzburg

In Cognitive Kin, Moral Strangers?, Judith Benz-Schwarzburg reveals the scope and relevance of cognitive kinship between humans and non-human animals. She presents a wide range of empirical studies on culture, language and theory of mind in animals and then leads us to ask why such complex socio-cognitive abilities in animals matter. Her focus is on ethical theory as well as on the practical ways in which we use animals. Are great apes maybe better described as non-human persons? Should we really use dolphins as entertainers or therapists? Benz-Schwarzburg demonstrates how much we know already about animals’ capabilities and needs and how this knowledge should inform the ways in which we treat animals in captivity and in the wild.


Edited by Aafke M.I. van Oppenraay

Aristotle's De Animalibus was an important source of zoological knowledge for the ancient Greeks and for medieval Arabs and Europeans. In the thirteenth century, the work was twice translated into Latin. One translation was produced directly from the Greek by William of Moerbeke. An earlier translation, made available as a critical edition in the present volume for the first time, was produced through an intermediary Arabic translation (Kitāb al-Ḥayawān) by Michael Scot (1175 - c. 1232). Scot's translation was one of the main sources of knowledge on animals in Europe and widely used until well into the fifteenth century. As a faithful translation of a translation produced by a Syriac-speaking Christian, the text contributes to our knowledge of Middle Arabic. The De Animalibus is composed of three sections: History of Animals (ten books), Parts of Animals (four books) and Generation of Animals (five books). Parts of Animals and Generation of Animals were published by BRILL as Volumes 5.2 and 5.3 of the book series ASL in 1998 (ASL 5.2) and 1992 (ASL 5.3). The present Volume 5.1.a contains the first section of Scot's translation of History of Animals: the general introduction and books 1-3, with Notes. Editions of the two concluding parts of History of Animals, ASL 5.1.b, books 4-6 and ASL 5.1.c, books 7-10, are in preparation. Complete Latin-Arabic and Arabic-Latin indices of History of Animals will be published in due course.

J. Alfredo Holley, Juliana Sterli and Néstor G. Basso

Pan-Chelidae (Testudines, Pleurodira) is a group of side-necked turtles with a currently disjointed distribution in South America and Australasia and characterized by two morphotypes: the long-necked and the short-necked chelids. Both geographic groups include both morphotypes, but different phylogenetic signals are obtained from morphological and molecular data, suggesting the monophyly of the long-necked chelids or the independent evolution of this trait in both groups. In this paper, we addressed this conflict by compiling and editing available molecular and morphological data for Pan-Chelidae, and performing phylogenetic and dating analyses over the individual and the combined datasets. Our total-evidence phylogenetic analysis recovered the clade Chelidae as monophyletic and as sister group of a clade of South American extinct chelids; furthermore Chelidae retained inside the classical molecular structure with the addition of extinct taxa in both the Australasian and the South American clades. Our dating results suggest a Middle Jurassic origin for the total clade Pan-Chelidae, an Early Cretaceous origin for Chelidae, a Late Cretaceous basal diversification of both geographic clades with the emergence of long-necked lineages, and an Eocene diversification at genera level, with the emergence of some species before the final breakup of Southern Gondwana and the remaining species after this event.

Gong Lin, Li Xinzheng and Lee Kwen-Shen

Two new species of genus Pheronemoides are described in this study. The Specimens were collected from the South China sea and the Caroline seamount in the northwestern Pacific Ocean. Pheronemoides crustiformis sp. nov. differs from its congeners in exhibiting large microamphidiscs, whip-like and slightly bent microuncinates and small spiny microdiactins. Pheronemoides curvipentactin sp. nov. possesses special pentactine atrialia with round terminal or tapering terminal curved pinular rays and crooked pentactins, making it easily distinguishable from its congeners. Partial sequences of the 28S rDNA and 16S rDNA genes were also amplified to confirm the family assignment of the two new species and to explore the systematic status of Pheronemoides.

Yutaro Oku, Kenji Iwao, Bert W. Hoeksema, Naoko Dewa, Hiroyuki Tachikawa, Tatsuki Koido and Hironobu Fukami

Recent molecular phylogenetic analyses of scleractinian corals have resulted in the discovery of cryptic lineages. To understand species diversity in corals, these lineages need to be taxonomically defined. In the present study, we report the discovery of a distinct lineage obscured by the traditional morphological variation of Fungia fungites. This taxon exists as two distinct morphs: attached and unattached. Molecular phylogenetic analyses using mitochondrial COI and nuclear ITS markers as well as morphological comparisons were performed to clarify their phylogenetic relationships and taxonomic positions. Molecular data revealed that F. fungites consists of two genetically distinct clades (A and B). Clade A is sister to a lineage including Danafungia scruposa and Halomitra pileus, while clade B formed an independent lineage genetically distant from these three species. The two morphs were also found to be included in both clades, although the attached morph was predominantly found in clade A. Morphologically, both clades were statistically different in density of septal dentation, septal number, and septal teeth shape. These results indicate that F. fungites as presently recognized is actually a species complex including at least two species. After checking type specimens, we conclude that specimens in clade A represent true F. fungites with two morphs (unattached and attached) and that all of those in clade B represent an unknown species and genus comprising an unattached morph with only one exception. These findings suggest that more unrecognized taxa with hitherto unnoticed morphological differences can be present among scleractinian corals.

Taeman Han, Seung-Hyun Kim, Hyung Joo Yoon, In Gyun Park and Haechul Park

The firefly subgenus Hotaria sensu lato of the genus Luciola currently includes four morphospecies: L. (H.) parvula, L. (H.) unmunsana, L (H.) papariensis, and L. (H.) tsushimana. The latter three are taxonomically controversial based on both morphological and molecular data. We examined the phylogenetic relationships and evolutionary history of the species and related congeners using partial COI gene sequences (DNA barcoding). Our phylogenetic analyses consistently supported the monophyly of Hotaria sensu lato, but did not resolve the generic rank. The two types of L. (H.) parvula in Japan can be considered distinct species that arose by pseudocryptic speciation during the Miocene, with substantial genetic divergence (15.41%). Three morphospecies, L. (H.) unmunsana, L (H.) papariensis, and L. (H.) tsushimana, split into several polyphyletic or paraphyletic groups, forming entangled species groups. They are considered an incipient group that is distinguishable genetically but not morphologically, with evidence for recent allopatric speciation events corresponding to geologic events and sea-level changes during the Pliocene and Pleistocene. Group III of L. (H.) unmunsana collected from the Jeolla region is a new taxon.

Laura Likov, Ante Vujić, Nataša Kočiš Tubić, Mihajla Đan, Nevena Veličković, Santos Rojo, Celeste Pérez-Bañón, Sanja Veselić, Anatolij Barkalov, Rüstem Hayat and Snežana Radenković

The putative monophyly and systematic position of Merodon nigritarsis group was assessed based on morphological and molecular data of the mitochondrial COI and nuclear 28S rRNA genes. The previously reported concept of the group has been redefined, and M. crassifemoris Paramonov, 1925 is now excluded. The related M. avidus group is redefined here, including the Merodon avidus complex and M. femoratus Sack, 1913. Species delimitation of morphologically defined species of M. nigritarsis group was well supported by COI gene analysis, with the exception of M. alagoezicus Paramonov, 1925 and M. lucasi Hurkmans, 1993. Descriptions are given for three new species of the M. nigritarsis species group: Merodon cohurnus Vujić, Likov et Radenković sp. n., Merodon longisetus Vujić, Radenković et Likov sp. n. and Merodon obstipus Vujić, Radenković et Likov sp. n., and one new species from the M. avidus group: Merodon rutitarsis Likov, Vujić et Radenković sp. n. A lectotype is designated for M. femoratus Sack, 1913, and two new synonymies of this species were proposed: M. biarcuatus Curran, 1939 and M. elegans Hurkmans, 1993. Here we review 18 species from the M. nigritarsis group and six species from the M. avidus group and provide morphological diagnoses of the species groups. Additionally, diagnosis of 12 branches (groups or individual taxa) of M. avidus-nigritarsis lineage, an illustrated diagnostic key for the males, and distribution map are provided for the new species.

Xin Tong and Bao-Zhen Hua

Neopanorpa, the second largest genus in Panorpidae, is mainly characterized by the well-developed notal organ on male tergum III. However, it remains largely unknown how the length of the notal organ influences the nuptial feeding behaviour of Neopanorpa. Here, we investigated the nuptial feeding by comparing the morphology of mating-related structures and the genital coupling of a) Neopanorpa lui Chou & Ran, 1981 with a weakly-developed notal organ, b) N. carpenteri Cheng, 1957 with a medium-sized notal organ, and c) N. longiprocessa Hua & Chou, 1997 with an extremely elongated notal organ. The couples of N. lui and N. carpenteri maintain an intermittent mouth-to-mouth mode but do not exchange any edible food. After that the males secrete a salivary mass onto the surface as a nuptial gift, which is distinctly larger in N. carpenteri than in N. lui. Correspondingly, the male salivary glands are more developed in N. carpenteri than in N. lui. Males of N. longiprocessa bear very short salivary glands corresponding to a coercive mating tactic. The genital couplings are similar among the three species of Neopanorpa. The paired hypovalves of males are used to control the cerci of females. The prominent basal processes of male gonostyli grasp the posterior portion of the female medigynium across the intersegmental membrane. The male aedeagus physically couples with the female medigynium to ensure the male phallotreme to connect to the female copulatory pore. The influence of the notal organ length on the nuptial feeding behaviour of Neopanorpa is briefly discussed.

Isabel T. Hyman and Frank Köhler

The helicarionid fauna of southeastern to mid-eastern Queensland is dominated by a group of semislugs with moderately reduced shells belonging to genera Fastosarion, Eungarion, Stanisicarion, Dimidarion, Macularion and Hymanarion. We comprehensively revise their systematic classification using comparative morpho-anatomy and mitochondrial phylogenetics, and demonstrate that these genera combined form a well-differentiated and monophyletic radiation. In our mitochondrial phylogeny, this radiation is divided into three main clades that are statistically well supported. One clade is also well defined in terms of diagnostic morpho-anatomical characters, but we could not identify diagnostic characters for the other two clades due to considerable levels of morpho-anatomical variation. We propose accepting only two genera, Fastosarion (with junior synonyms Eungarion, Dimidarion, and Hymanarion) and Stanisicarion (with junior synonym Macularion). Both genera represent mutually monophleytic sister taxa that can consistently be distinguished by the presence or absence of a penial verge that is fused to the penial wall and by egg shape. We also synonymise Fastosarion ameyi with F. aquavitae, F. schelli with F. helenkingae, Dimidarion peterbrocki and D. slatyeri with F. alyssa, Stanisicarion virens with S. freycineti. Revised species descriptions are presented for Fastosarion alyssa, F. aquavitae, F. brazieri, F. comerfordae, F. griseolus, F. hannianus, F. helenkingae, F. mcdonaldi, F. minerva, F. paluma, F. papillosus, F. pustulosus, F. superbus, Stanisicarion aquila and S. freycineti. Nine new species, Fastosarion deensis, F. ephelis, F. insularis, F. katatonos, F. longimentula, F. rowani, F. sarina, F. tuljun and Stanisicarion wolvi are described, bringing the total number of accepted species to 24.

Somayeh Esmaeili-Rineh, Mahmoud Mamaghani-Shishvan, Cene Fišer, Vahid Akmali and Nargess Najafi

The connectivity of groundwater aquifers is lower compared to surface waters. Consequently, groundwater species are expected to have smaller distributional ranges than their surface relatives. Molecular taxonomy, however, unveiled that many species comprise complexes of morphologically cryptic species, with geographically restricted distributional ranges in subterranean as well as in surface waters. Hence, the range sizes of surface and groundwater species might be more similar in size than hitherto thought. We tested this hypothesis by comparing the range size of surface amphipods of the genus Gammarus and subterranean amphipods of the genus Niphargus in Iran. We re-analyzed the taxonomic structure of both genera using two unilocus species delimitation methods applied to a fragment of the COI mitochondrial marker, to identify molecular operational taxonomic units (MOTUs), and assessed the maximum linear extent (MLE) of the ranges of MOTUs from both genera. Genus Gammarus comprised 44–58 MOTUs while genus Niphargus comprised 20–22 MOTUs. The MLEs of the two genera were not significantly different, regardless the delimitation method applied. The results remained unchanged also after exclusion of single site MOTUs. We tentatively conclude that in this case there is no evidence to consider that groundwater species are geographically more restricted than surface species.

Giulia Perina, Ana I. Camacho, Joel Huey, Pierre Horwitz and Annette Koenders

The stygofaunal family of Bathynellidae, is an excellent group to study the processes that shape diversity and distribution, since they have unknown surface or marine relatives, high level of endemism, and limited dispersal abilities. Recent research on Bathynellidae in Western Australia (Pilbara) has uncovered new taxa with unexpected distributions and phylogenetic relationships, but the biogeographical processes that drive their diversification on the continent are still unclear. By exploring the diversity, distribution, and divergence time of Bathynellidae in a setting such as the perched and isolated aquifers of the Cleaverville Formation in the north of the De Grey River catchment (Pilbara), we aim to test the hypothesis that vicariance has shaped the distribution of this family, specifically if one or multiple vicariant events were involved. We analysed the specimens collected from perched water in different plateaus of the Cleaverville Formation, combining morphological and molecular data from mitochondrial and nuclear genes. We described two new species and genera (Anguillanella callawaensis gen. et sp. nov. and Muccanella cundalinensis gen. et sp. nov.), and two additional taxa are recognised using morphology and/or Automatic Barcode Gap Discovery and Poisson Tree Processes species delimitation methods. New genera and species result restricted to isolate perched aquifers on single plateaus and their distributions, phylogenetic relationships, and divergence time estimates support multiple vicariant events and ancient allopatric speciation.

Javier H. Santos-Santos, Mireia Guinovart-Castán and David R. Vieites

Mantellid frogs present an extensive adaptive radiation endemic to Madagascar and Comoros, being the subfamily Mantellinae the most morphologically and ecologically diverse. The Mantellinae present key innovative evolutionary traits linked to their unique reproductive behavior, including the presence of femoral glands and a derived vomeronasal organ. In addition, previous studies pointed to size differentiation in playing an important role in species’ dispersal capacities and shaping of their geographic ranges. Despite the high phenotypic variation observed in this clade, to date an exhaustive morphological analysis of their anatomy has still not been performed, much less in relation to internal structures. Here, we present a comprehensive skeletal description of a mantellid species, Blommersia transmarina, from the island of Mayotte in the Indian Ocean, which has potentially undergone a process of moderate gigantism compared to other Blommersia species. We describe its intraspecific skeletal variation utilizing non-destructive volume renderings from μCT-scans, and characterize the presence of sexual dimorphism and size covariation in skeletal structures. Notably, we found numerous signs of hyperossification, a novel structure for mantellids: the clavicular process, and the presence of several appendicular sesamoids. Our findings suggest that skeletal phenotypic variation in this genus may be linked to biomechanical function for reproduction and locomotion.

Maliheh Pirayesh Shirazinejad, Mansour Aliabadian and Omid Mirshamsi

The white wagtail (Motacilla alba) species complex with its distinctive plumage in separate geographical areas can serve as a model to test evolutionary hypotheses. Its extensive variety in plumage, despite the genetic similarity between taxa, and the evolutionary events connected to this variety are poorly understood. Therefore we sampled in the breeding range of the white wagtail: 338 individuals were analyzed from 74 areas in the Palearctic and Mediterranean. We studied the white wagtail complex based on two mitochondrial DNA markers to make inferences about the evolutionary history. Our phylogenetic trees highlight mtDNA sequences (ND2, CR), and one nuclear marker (CHD1Z), which partly correspond to earlier described clades: the northern Palearctic (clade N); eastern and central Asia (clade SE); south-western Asia west to the British Isles (clade SW); and Morocco (clade M). The divergence of all clades occurred during the Pleistocene. We also used ecological niche modelling for three genetic lineages (excluding clade M); results showed congruence between niche and phylogenetic divergence in these clades. The results of the white wagtail ancestral area reconstruction showed the influence of dispersal on the distribution and divergence of this complex species. The most important vicariance event for the white wagtail complex may have been caused by the Gobi and Taklamakan deserts. We conclude that the ancestral area of the white wagtail complex was probably in the Mediterranean, with its geography having a considerable effect on speciation processes.

Oksana A. Korzhavina, Bert W. Hoeksema and Viatcheslav N. Ivanenko

This review of copepod crustaceans associated with reef-dwelling cnidarians, sponges and echinoderms of the Greater Caribbean is based on published records, systematically arranged by the classification of symbiotic copepods and their hosts, sampling sites, coordinates, depth and date of sampling, literature sources, and three recent surveys (Cuba, St. Eustatius in the Eastern Caribbean and Curaçao in the Southern Caribbean). This resulted in totals of 532 records of 115 species of symbiotic copepods (47 genera, 17 families, three orders) hosted by 80 species of invertebrates, representing scleractinians (47%), octocorals (9%), echinoderms (3%), and sponges (1%). Among ten Caribbean ecoregions, the Greater Antilles (with 64 species of symbiotic copepods) as well as the Southern and Eastern Caribbean (with 46 and 17 species of copepods, respectively) are the most studied and best represented, whereas only six species of copepods are known from Bermuda, one from Southwestern Caribbean and none from the Gulf of Mexico. The absence of poecilostomatoid copepods (Anchimolgidae, Rhynchomolgidae and Xarifidae) on Caribbean stony corals as noted by Stock (1988) is confirmed. The results indicate that the diversity and ecology of Caribbean symbiotic copepods are still poorly investigated.

Tomáš Němec and Michal Horsák

Shell formation is the main defensive strategy against predation for the majority of snails. Therefore, various predators have had to develop a variety of techniques how to overcome this barrier. As shells can persist in a calcium-rich environment for a long time, specific external or internal traces on shells left by predators indicate whether and who killed the snail. Based on litter samples collected at 30 sites of five different habitat types, the intensity and type of predation were assessed. The minimal predation rate varied between 0.0 and 21%, with an average of 8%. The highest rate was observed at limestone steppes, on average 15%. Beetles were found to be the most common predators of snails; however, predation by snails was more common in calcareous fens. Predation by some vertebrates and dipteran flies was also recognised. To test the role of mouth barriers as a means to reduce predation by carabid beetles that break the shell from an aperture, we analysed the predation rate separately on adult and juvenile shells using 24 populations of the steppe snail Granaria frumentum (Draparnaud, 1801). As expected, carabid beetles chiefly preferred juveniles compared to adult shells (Wilcoxon test, p < 0.001). On the contrary, the parasitoid fly Pherbellia limbata (Meigen, 1830) and Drilus beetles preferred adults. We found that predation by carabid beetles positively increased with prey abundance (R2 = 42.8%, p = 0.021), while no relation was observed for the parasitoid (p = 0.703), likely due to their feeding specialisation.

Miguel A. Meca, Pilar Drake and Daniel Martin

The polychaete Oxydromus okupa lives in association with the bivalves Scrobicularia plana and Macomopsis pellucida in the intertidal of Río San Pedro (CI = Cádiz Intertidal) and adjacent to CHipiona (CH) harbour, and in the subtidal of the Bay of Cádiz (CS = Cádiz Subtidal). We analyse these populations morphometrically, ecologically (including infestation characteristics) and genetically (intertidal populations, 16S and ITS-1 genes). We consider “host”, “environment” and the combined “host and environment” as possible factors of interpopulation variability. Morphometry revealed three well-defined clusters for CI, CH and CS, showing intergroup phenotypic differences ranging from 35 to 50%. Hosts shell lengths ranged between 26 and 36 mm for S. plana and 20 and 28 mm for M. pellucida. The infestation of small M. pellucida by juvenile O. okupa suggests they show an active size segregation behaviour. The intertidal seems to be less favourable (infestation rate <25% vs. up to 65% in the subtidal), and did not show recent bottleneck events. Overall, CI and CH were genetically homogeneous, but showed a significant divergence (one dominant haplotype in each host species), suggesting host shift as being a soft barrier to gene flow. Most characters related with host-entering varied among populations, suggesting symbiotic behaviour to play a key role in reducing panmixia and leading to the initial phases of a speciation process in sympatric symbiotic populations. Polyxeny and symbiotic behaviour in O. okupa seem thus to be underlying mechanisms contributing to its great phenotypic variety, marked ecological differences, and genetic divergence.

Giacinta Angela Stocchino, Ronald Sluys, Abdel Halim Harrath, Lamjed Mansour and Renata Manconi

Invasions of alien species form one of the major threats to global biodiversity. Among planarian flatworms many species are known to be invasive, in several cases strongly affecting local ecosystems. Therefore, a detailed knowledge on the biology of an invasive species is of utmost importance for understanding the process of invasion, the cause of its success, and the subsequent ecological impact on native species. This paper provides new information on the biology of introduced populations of the freshwater flatworm Girardia tigrina (Girard, 1850) from Europe. This species is a native of the Nearctic Region that was accidentally introduced into Europe in the 1920s. Since then, numerous records across the European continent bear witness of the invasiveness of this species, although only a few studies focused on the biology of the introduced populations. We report on the morphology of sexualized individuals from a fissiparous Italian population, representing the second record of spontaneous sexualization of fissiparous individuals in this species. A detailed morphological account of the reproductive apparatus of these ex-fissiparous animals is presented. Our results increased the number of morphological groups previously recognized for European populations of G. tigrina, thus corroborating the hypothesis on multiple independent introductions to this continent. Karyological results obtained from our fissiparous Italian individuals revealed a constant diploid chromosome complement of sixteen chromosomes. Further, we document the marked intraspecific variation in several morphological features of this species.

Cessa Rauch, Bert W. Hoeksema, Bambang Hermanto and Charles H.J.M. Fransen

Most marine palaemonid shrimp species live in symbiosis with invertebrates of various phyla. These associations range from weak epibiosis to obligatory endosymbiosis and from restricted commensalism to semi-parasitism. On coral reefs, such symbiotic shrimps can contribute to the associated biodiversity of reef corals. Among the host taxa, mushroom corals (Cnidaria: Anthozoa: Fungiidae) are known to harbour various groups of symbionts, including shrimps. Some but not all of these associated species are host-specific. Because data on the host specificity of shrimps on mushroom corals are scarce, shrimp species of the genus Periclimenes were collected from mushroom corals during fieldwork in Lembeh Strait, North Sulawesi, Indonesia. Using molecular (COI barcoding gene) and morphological methods, three species of Periclimenes were identified: P. diversipes, P. watamuae and a species new to science, P. subcorallum sp. nov., described herein. Their host specificity was variable, with eight, three and two fungiid host records, respectively. It is concluded that shrimp species of the genus Periclimenes show much overlap in their host choice and that particular morphological traits in the host species appear to play a more important role than phylogenetic affinities within the host group.

Gerrit Potkamp and Charles H.J.M. Fransen

Over the last century, a large body of literature emerged on mechanisms driving speciation. Most of the research into these questions focussed on terrestrial systems, while research in marine systems lagged behind. Here, we review the population genetic mechanisms and geographic context of 33 potential cases of speciation with gene flow in the marine realm, using six criteria inferred from theoretical models of speciation. Speciation with gene flow occurs in a wide range of marine taxa. Single traits, which induce assortative mating and are subjected to disruptive selection, such as differences in host-associations in invertebrates or colour pattern in tropical fish, are potentially responsible for a decrease in gene flow and may be driving divergence in the majority of cases. However, much remains unknown, and with the current knowledge, the frequency of ecological speciation with gene flow in marine systems remains difficult to estimate. Standardized, generally applicable statistical methods, explicitly testing different hypotheses of speciation, are, going forward, required to confidently infer speciation with gene flow.

Ennominae II

(Boarmiini, Gnophini, additions to previous volumes)


Bernd Müller, Sven Erlacher, Axel Hausmann, Hossein Rajaei, Pasi Sihvonen and Peder Skou

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Final part of The Geometrid Moths of Europe concluding the revision of the European Ennominae moths, covering a total of 181 species plus 21 species of Geometridae found in Europe since publication of previous volumes. Several difficult genera such as Charissa, Psodos, Sciadia, Nychiodes, Selidosema, Peribatodes and the Tephronia complex are covered. Four new species are described, and the fauna is richly illustrated by 1116 specimens in 30 colour plates, 131 genitalia plates and numerous text-figures highlighting diagnostic features. For each species a taxonomic summary, description and diagnosis, distribution map, biology and genetic data are provided. Over 140 taxonomic changes are proposed. A systematic, annotated checklist with synonyms is provided, which summarises the entire Geometridae fauna of Europe (999 species) and adjacent regions.

Piotr Gąsiorek, Daniel Stec, Witold Morek and Łukasz Michalczyk

Isohypsibioidea are most likely the most basally branching evolutionary lineage of eutardigrades. Despite being second largest eutardigrade order, phylogenetic relationships and systematics within this group remain largely unresolved. Broad taxon sampling, especially within one of the most speciose tardigrade genera, Isohypsibius Thulin, 1928, and application of both comparative morphological methods (light contrast and scanning electron microscopy imaging of external morphology and buccal apparatuses) and phylogenetic framework (18S + 28S rRNA sequences) resulted in the most comprehensive study devoted to this order so far. Two new families are erected from the currently recognised family Isohypsibiidae: Doryphoribiidae fam. nov., comprising all aquatic isohypsibioids and some terrestrial isohypsibioid taxa equipped with the ventral lamina; and Halobiotidae fam. nov., secondarily marine eutardigrades with unique adaptations to sea environment. We also split Isohypsibius into four genera to accommodate phylogenetic, morphological and ecological variation within the genus: terrestrial Isohypsibius s.s. (Isohypsibiidae), with smooth or sculptured cuticle but without gibbosities; terrestrial Dianea gen. nov. (Isohypsibiidae), with small and pointy gibbosities; terrestrial Ursulinius gen. nov. (Isohypsibiidae), with large and rounded gibbosities; and aquatic Grevenius gen. nov. (Doryphoribiidae fam. nov.), typically with rough cuticle and claws with branches of very similar heigths. Claw morphology is reviewed and, for the first time, shown to encompass a number of morphotypes that correlate with clades recovered in the molecular analysis. The anatomy of pharynx and cuticle are also shown to be of high value in distinguishing supraspecific taxa in Isohypsibioidea. Taxonomy of all isohypsibioid families and genera is discussed, with special emphasis on the newly erected entities. Finally, a dychotomous diagnostic key to all currently recognised isohypsibioid families and genera is provided.

Bernhard A. Huber and Anne Chao

Ratio-like approaches for estimating global species richness have been criticised for their unjustified extrapolation from regional to global patterns. Here we explore the use of cumulative percentages of ‘new’ (i.e., not formally described) species over large geographic areas (‘megatransects’) as a means to overcome this problem. In addition, we take into account undetected species and illustrate these combined methods by applying them to a family of spiders (Pholcidae) that currently contains some 1,700 described species. The raw global cumulative percentage of new species (‘new’ as of the end of 2008, when 1,001 species were formally described) is 75.1%, and is relatively constant across large biogeographic regions. Undetected species are estimated using the Chao2 estimator based on species incidence data (date by species and locality by species matrices). The estimated percentage of new species based on the date by species matrices is 76.0% with an estimated standard error (s.e.) of 2.6%. This leads to an estimated global species richness of about 4,200 with a 95% confidence interval of (3,300, 5,000). The corresponding values based on locality by species matrices are 84.2% (s.e. 3.0%) and 6,300 with a 95% confidence interval of (4,000, 8,600). Our results suggest that the currently known 1,700 species of Pholcidae may represent no more than about 25–40% of the total species richness. The impact of further biasing factors like geography, species size and distribution, cryptic species, and model assumptions needs to be explored.

Antonio Archidona-Yuste, Carolina Cantalapiedra-Navarrete, Pablo Castillo and Juan E. Palomares-Rius

The genus Longidorus constitutes a large group of approximately 170 species of plant-ectoparasitic nematodes that are polyphagous and distributed almost worldwide. Some of the species of this genus are vectors of plant viruses. Species discrimination in Longidorus is difficult because the morphology is very conservative, and morphometric characters often overlap, leading to potential misidentification. Integrative taxonomy, based on the combination of molecular analyses with morphology, is a useful and necessary approach in Longidorus species identification. In Spain from 2014 to 2017, we conducted nematode surveys among cultivated and wild plants, from which we identified 13 populations of Longidorus, two of which appeared to represent new species and are described herein as L. iliturgiensis sp. nov. and L. pacensis sp. nov., and 11 populations belonging to eight known species: L. africanus, L. baeticus, L. carpetanensis, L. fasciatus, L. nevesi, L. cf. olegi, L. pini, and L. vallensis. Three species are new geographical records for Spain (L. nevesi, L. cf. olegi, and L. africanus). We report molecular data for L. nevesi, L. cf. olegi, L. carpetanensis and L. pini for the first time. Additionally, we describe the males of L. pini and the juveniles of L. cf. olegi.

Yee Wah Lau, Frank R. Stokvis, Yukimitsu Imahara and James D. Reimer

Stoloniferan octocorals (Cnidaria: Anthozoa: Octocorallia: Alcyonacea) are a relatively unexplored fauna in the Ryukyus (southern Japan), known to be a tropical marine region of high biodiversity and endemism of species. Specimens of stoloniferous octocorals were collected during fieldwork along the coasts of two islands (Okinawa and Iriomote) in the Okinawa Prefecture. Despite their phenotypic polyp variation, this study shows their morphological and molecular uniqueness, leading to the description of a new genus with a single species: Hanabira yukibana, gen. nov., sp. nov. They are placed within the Clavulariidae and form a sister clade basally to the genus Knopia Alderslade & McFadden, 2007 and species of Clavularia Blainville, 1830. The polyps of this new species show morphological variation in both shape and sclerite density, but there is conformity in the typical overall petal shaped tentacles, which have fused pinnules (pseudopinnules). Depending on the densities of their sclerites and their photosynthetic endosymbiotic algae (zooxanthellae) of the family Symbiodiniaceae, there is a characteristic sheen present in the tentacles. Moreover, the zooxanthellae hosted by our specimens form a clear, small-scale biogeographic pattern; all H. yukibana specimens from Okinawa Island contained zooxanthellae of the genus Cladocopium Lajeunesse & H.J. Jeong, 2018 (= former Symbiodinium ‘Clade C’) and all specimens from Iriomote Island hosted zooxanthellae of the genus Durusdinium LaJeunesse, 2018 (= former Symbiodinium ‘Clade D’). These results show the potential for variation among the Symbiodiniaceae floras within octocorals, something that has not yet been investigated for the large majority of zooxanthellate octocoral species.

Tineidae II

(Myrmecozelinae, Perissomasticinae, Tineinae, Hieroxestinae, Teichobiinae and Stathmopolitinae)


Reinhard Gaedike

This second volume on Tineidae treats the subfamilies Myrmecozelinae, Perissomasticinae, Tineinae, Hieroxestinae, Teichobiinae and Stathmopolitinae of Europe. It presents information for the identification of 103 species of tineid moths. Information is added on the life history and distribution of each species. The distribution data are summarized in a table showing the records for each European country. 23 scientific names are synonymized and two taxa previously regarded as synonyms have proved to represent valid species.
Additional records are listed for species treated in volume 7, as well as two taxa which were overlooked before and nine new species are listed.


Angelica Groom

The book examines the roles that rare and exotic animals played in the cultural self-fashioning and the political imaging of the Medici court during the family’s reign, first as Dukes of Florence (1532-1569) and subsequently as Grand Dukes of Tuscany (1569-1737). The book opens with an examination of global practices in zoological collecting and cultural uses of animals. The Medici’s activities as collectors of exotic species, the menageries they established and their deployment of animals in the ceremonial life of the court and in their art are examined in relation to this wider global perspective. The book seeks to nuance the myth promoted by the Medici themselves that theirs was the most successful princely serraglio in early modern Europe.  

Neil Cumberlidge and Savel R. Daniels

Edited by R. Vonk

Surveys of the freshwater crabs of two islands in the Gulf of Guinea, Central Africa, allowed a revision of the taxonomy of two little-known island endemic species, based for the first time on adult males: Potamonautes margaritarius (A. Milne-Edwards, 1869) from São Tomé, and of P. principe Cumberlidge, Clark and Baillie, 2002, from Príncipe (Brachyura; Potamonautidae). A new species of Potamonautes from southern São Tomé (Potamonautes saotome sp. nov.) is also described that is genetically distinct and has a clearly separate geographic distribution from P. margaritarius from northern São Tomé. The new species from southern São Tomé can be recognized by a suite of characters of the carapace, thoracic sternum, and gonopods. The taxonomy of P. margaritarius (A. Milne-Edwards, 1869) is stabilized by selecting a neotype from northern São Tomé. Potamonautes principe from Príncipe is the most distinct of the three taxa, with a more swollen carapace that has smooth anterolateral margins, and a shorter, straighter male first gonopod. All three taxa are morphologically distinct species that have also been clearly distinguished as evolutionarily separate lineages by mtDNA analysis and haplotyping in an earlier study. Previous phylogenetic evidence supports two separate island colonization events at different times in the past from different ancestral populations, one to São Tomé and another to Príncipe that resulted in the establishment of the endemic freshwater crab faunas of these two islands.

Sharon E. Kessler, Ute Radespiel, Alida I. F. Hasiniaina, Leanne T. Nash and Elke Zimmermann

Edited by V. Nijman

Frequent kin-biased coalitionary behaviour is a hallmark of mammalian social complexity. Furthermore, selection to understand complex social dynamics is believed to underlie the co-evolution of social complexity and large brains. Vocalisations have been shown to be an important mechanism with which large-brained mammals living in complex social groups recognise and recruit kin for coalitionary support during agonistic conflicts. We test whether kin recognition via agonistic calls occurs in a small-brained solitary foraging primate living in a dispersed social network, the grey mouse lemur (Microcebus murinus, Miller JF, 1777). As mouse lemurs are frequent models for ancestral solitary foraging mammals, this study examines whether kin recognition via agonistic calls could be the foundation from which more complex, kin-based coalitionary behaviour evolved. We test whether female wild mouse lemurs in Ankarafantsika National Park, Madagascar, react differently to agonistic calls from kin and nonkin and to calls from familiar and unfamiliar individuals during playback experiments. Subjects showed no significant differences in reactions to the different stimuli; thus they did not react differently based upon kinship or familiarity. Results suggest that this solitary foraging species does not use agonistic calls to recognise kin and monitor agonistic interactions involving kin, unlike several species of Old World monkeys and hyenas. Thus, kin recognition via agonistic calls may have evolved independently in these lineages in parallel with greater social complexity and frequent coalitionary behaviour.

Alejandro Sánchez-Chardi, Marian García-Pando, Eudald Pujol-Buxó, Maria A. Sans-Fuentes, María José López-Fuster and Francesc Muñoz-Muñoz

Edited by A. Ivanović

Island populations of terrestrial mammals often undergo extensive behavioural and morphological changes when separated from mainland populations. Within small mammals these changes have been mainly reported in rodents but were poorly assessed in soricomorphs. In this study we compared mandible morphology and body condition between mainland and island populations of the greater white-toothed shrew, Crocidura russula. The results indicated that island specimens were bigger and heavier than the mainland counterpart, and they showed changes in mandible shape that were associated with higher mechanical potentials. We suggest that these changes might be the result of the interaction of two main factors taking place in the island population: ecological release (i.e. the decrease of predation and interspecific competition), and consequently the increase of intraspecific competition. While the increase in size and body condition in island shrews could be a direct result from reduced predation and interspecific competition, the changes in mandible shape and the increase of both mechanical potential and sexual dimorphism could have arisen indirectly as a response to stronger intraspecific competition.

Ana I. Camacho, Paloma Mas-Peinado, Santi Watiroyram, Anton Brancelj, Elia Bandari, Beatriz A. Dorda, Adrián Casado and Isabel Rey

Edited by R. Vonk

A preliminary molecular phylogenetic framework for 12 genera (23 species) of the family Parabathynellidae from Europe, Australia, North Africa and India is presented based on mitochondrial and nuclear markers (Cox1 and 18S). The generated hypothesis places the Southeast Asia genus Paraeobathynella closer to European genera (Iberobathynella, Paraiberobathynella and Parabathynella) than to the Australian (Brevisomabathynella, Atopobathynella, Billibathynella, Octobathynella, Arkaroolabathynella and Lockyerenella) or Indian genera (Habrobathynella), or to the cosmopolitan genus Hexabathynella (Spain and Australia). Three new species of the genus Paraeobathynella from Thailand, P. ratensis n. sp., P. siamensis n. sp. and P. hanjavanitiana n. sp., are described based on morphological and molecular features. This is the first record of the genus from Thailand and extends its range of distribution within Asia, where it was previously known only from Vietnam. The new species are clearly separated as independent units at least since the Middle Miocene.

Antonia Chroni, Ana Grković, Jelena Ačanski, Ante Vujić, Snežana Radenković, Nevena Veličković, Mihajla Djan and Theodora Petanidou

Edited by A. Ivanović

This study provides an overview of the Eumerus minotaurus taxon group, diagnosing a new species, E. anatolicus Grković, Vujić and Radenković sp. n. (Muğla, Turkey), and unraveling three cryptic species within E. minotaurus: E. karyates Chroni, Grković and Vujić sp. n. (Peloponnese, Greece), E. minotaurus Claussen and Lucas, 1988 (Crete and Karpathos, Greece) and E. phaeacus Chroni, Grković and Vujić sp. n. (Corfu and Mt Olympus, Greece; Mt Rumija, Montenegro). We applied an integrative taxonomic approach based on molecular, morphological and wing geometric morphometric data to corroborate and delimit cryptic species within the complex. In addition, we discuss the latent biogeographic patterns and speciation processes leading to configuration of the E. minotaurus group based on palaeogeographic evolution of the Aegean. Mitochondrial phylogeographic analysis suggested that speciation within the E. minotaurus group is attributable to formation of the mid-Aegean Trench and Messinian Salinity Crisis, and was integrated at the Pleistocene. We show that more accurate estimates of divergence times may be based on geological events rather than the standard arthropod mtDNA substitution rate.

Field Guide to the Birds of Suriname

Revised and Updated Second Edition


Arie L. Spaans, Otte H. Ottema and Jan Hein J.M. Ribot

Suriname, located on the Atlantic coast of northeastern South America, is a relatively small country compared to most other South American countries. It nevertheless has a rich avifauna. By the end of 2017, 751 species (including 765 subspecies) were known to occur in Suriname. Most of the land area of Suriname is still covered with tropical rainforest and the country should be a must-visit for birdwatchers. Suriname is even mentioned as being the best country to spot certain neotropical species. Surprisingly, few birders visit Suriname. The main reason given is the lack of a handy pocket guide that can easily be carried in a backpack. This (revised and updated) edition of the Field Guide to the Birds of Suriname (with its 109 color plates) tries to fill this gap. In addition to species accounts, data on topography, climate, geology, geomorphology, biogeography, avifauna composition, conservation, and hotspots for bird watching are given. So, why delay your trip to this beautiful and friendly country any longer? Suriname with its rich avifauna awaits you!

Humberto G. Ferrón, María Victoria Paredes-Aliaga, Carlos Martínez-Pérez and Héctor Botella

Edited by M. Rücklin

Galeomorph sharks constitute the most taxonomically and ecologically diverse superorder of living selachians. Despite comprising several typically deep-water taxa, no bioluminescent species have been reported in this group so far. Interestingly, the study of shark squamation has been revealed in recent years to be a good proxy for inferring some ecological aspects of poorly known species. In particular, the high morphological specificity of the dermal denticles and the squamation patterns of all currently-known bioluminescent sharks could constitute a potential tool for predicting bioluminescence in both fossil and living taxa. Following this idea, we provide the first evidence supporting the possible existence of bioluminescence among galeomorph sharks by means of the quantitative study of Apristurus ampliceps squamation pattern. Classical morphometric analysis and multivariate statistical procedures have allowed us to determine that A. ampliceps squamation, composed mainly of bristle-shaped dermal denticles, is highly convergent with that of the bioluminescent shark Etmopterus spinax. The ecology of A. ampliceps, being a species that exclusively inhabits aphotic waters, is in agreement with such a morphofunctional interpretation, but finding photophores is imperative to confirm this prediction.

Jan W. Arntzen

Edited by A. Ivanović

I document the contiguous distribution of the marbled newt species Triturus marmoratus and T. pygmaeus over the western part of the Iberian peninsula with a suite of morphological and molecular genetic data from altogether 141 populations. Morphological characters that identify the species are body size and the colour pattern character ‘Links’. Links is the number of transversal connections following the green surface at the lateral sides of the newts’ body. Large adults with few Links are T. marmoratus and small adults with many Links are T. pygmaeus. However, no morphological identification criterion is entirely adequate. Eight molecular genetic markers show markedly bimodal character state distributions that give rise to sharp species range descriptions, with T. marmoratus in the north of the Iberian peninsula and T. pygmaeus in the south and along most of the Atlantic coast of Portugal. I encountered ten genetically admixed populations that are all located at the T. marmoratus - T. pygmaeus species range interface, suggesting widespread but limited interspecific hybridization. A latitudinal transect across Portugal confirmed the narrow and steep transition from one to the other species for morphological and molecular characters alike. In central Portugal the position of the hybrid zone coincides with the river Tejo. However, the cline for mitochondrial DNA is relatively wide and shallow and its centre is positioned south of the river. In view of published data that reconstruct the northward advance of T. pygmaeus along the Portuguese coast at the expense of T. marmoratus, I propose that T. marmoratus had a wider range in central Portugal too, where it was eventually superseded by T. pygmaeus. I hypothesize that ‘marmoratus’ mtDNA haplotypes found south of the Tejo constitute a ‘genetic footprint’ left behind in T. pygmaeus by the receding species T. marmoratus.

Jia Jin Marc Chang, Ywee Chieh Tay, Hui Ping Ang, Karenne Phyu Phyu Tun, Loke Ming Chou, Rudolf Meier and Danwei Huang

Edited by B.W. Hoeksema

Recent years have seen an increase in the number of studies that use DNA sequence information in addition to morphological methods, as the latter alone can be inadequate for morphologically similar, cryptic species. Marine onchidiid slugs (Mollusca: Gastropoda: Onchidiidae), which commonly inhabit intertidal environments and have a near-global distribution, comprise 11 genera and 86 described species. Singapore has 19 recorded species of onchidiids with Peronia verruculata (Cuvier, 1830) being the most abundant. Here we present mitochondrial DNA sequence data from 63 P. verruculata-like specimens to show that they constitute two closely-related clades with distinct sequence signatures and a clear barcode gap. Intraclade distances are 0–1.8% while interclade distances range from 4.7 to 6.0%. Scanning electron microscopy of internal copulation organs reveals that the two clades have distinct penial and accessory gland spine structures, which also differ in size. Using an integrative taxonomic approach, we propose that what was initially thought to be P. verruculata in Singapore, actually are two different species of Peronia.

Vukica Vujić, Bojan Ilić, Zvezdana Jovanović, Sofija Pavković-Lučić, Sara Selaković, Vladimir Tomić and Luka Lučić

Edited by A. Minelli

Sexual selection can be a major driving force that favours morphological evolution at the intraspecific level. According to the sexual selection theory, morphological variation may accompany non-random mating or fertilization. Here both variation of linear measurements and variation in the shape of certain structures can significantly influence mate choice in different organisms. In the present work, we quantified sexual behaviour of the millipede Megaphyllum bosniense (Verhoeff, 1897) as characterized by several sequences. These are: mating latency, duration of copulation, contact to copulation time, duration of contact without copulation, time from entrance (time-point when individuals were placed in boxes in which tests occurred) to contact with copulation, and time from entrance to contact without copulation. Further, we analysed the influence of morphological variation (both variation of linear measurements and variation in the shape of several structures) on mating success. Variation of body length, antennal length, length of the walking legs, trunk width, and trunk height was analysed by traditional morphometrics, while variation in size and shape of the antennae, walking legs, head, and gonopods (promeres, opisthomeres) was analysed using geometric morphometrics. More than half of all physical contacts detected among the millipedes resulted in copulation. Based on the value of sexual selection coefficients, preferences toward the previous partner were found to be prevalent in both female and male choice tests. Individuals with different mating status significantly differed in some morphological traits (body mass, head centroid size, head shape, and promere shape). Our study yielded new information about the sexual behaviour of millipedes and variation of morphological traits as a potential basis for mate preferences.

Nikol Kmentová, Maarten Van Steenberge, Joost A.M. Raeymaekers, Stephan Koblmüller, Pascal I. Hablützel, Fidel Muterezi Bukinga, Théophile Mulimbwa N’sibula, Pascal Masilya Mulungula, Benoît Nzigidahera †, Gaspard Ntakimazi, Milan Gelnar and Maarten P.M. Vanhove

Edited by R. Sluys

Whereas Lake Tanganyika’s littoral and benthic zones are famous for their diverse fish communities, its pelagic zone is dominated by few species, of which two representatives of Clupeidae (Limnothrissa miodon and Stolothrissa tanganicae) take a pivotal role. We investigated the monogenean fauna infecting these freshwater clupeids to explore the link between parasite morphology and host species identity, or seasonal and geographical origin, which may reveal host population structure. Furthermore, we conducted phylogenetic analyses to test whether these parasitic flatworms mirror their host species’ marine origin. Based on 406 parasite specimens infecting 385 host specimens, two monogenean species of Kapentagyrus Kmentová, Gelnar and Vanhove, gen. nov. were morphologically identified and placed in the phylogeny of Dactylogyridae using three molecular markers. One of the species, Kapentagyrus limnotrissae comb. nov., is host-specific to L. miodon while its congener, which is new to science and described as Kapentagyrus tanganicanus Kmentová, Gelnar and Vanhove, sp. nov., is infecting both clupeid species. Morphometrics of the parasites’ hard parts showed intra-specific variability, related to host species identity and seasonality in K. tanganicanus. Significant intra-specific differences in haptor morphometrics between the northern and southern end of Lake Tanganyika were found, and support the potential use of monogeneans as tags for host population structure. Based on phylogenetic inference, we suggest a freshwater origin of the currently known monogenean species infecting clupeids in Africa, with the two species from Lake Tanganyika representing a quite distinct lineage.

Davide Maggioni, Roberto Arrigoni, Paolo Galli, Michael L. Berumen, Davide Seveso and Simone Montano

Edited by B.W. Hoeksema

The Zancleidae is a hydrozoan family that currently comprises three genera and 42 nominal species. The validity of numerous taxa in this family still needs to be assessed with integrative analyses and complete life cycle descriptions. The vast majority of its species live symbiotically with other organisms, among which cheilostomate bryozoans are the most common hosts. These bryozoan-associated zancleids are host-specific and encompass all species of the genera Halocoryne and Zanclella, as well as several species in the genus Zanclea. Zancleids show variable morphologies, including highly reduced polyps and medusae. Their phylogenetic history is uncertain due to the often intergrading morphologies and the shortage of molecular data. In the present study, two species of Zanclea from the Indian Ocean and the Red Sea are analysed, using morphological and DNA-based approaches. Morphological analyses of the polyp and medusa stages show that, despite a general resemblance with each other and with Zanclella diabolica, the two species differ in some characters, and this is supported also by molecular investigations. The DNA analyses show that the two species are monophyletic and closely related, but divergent from other Zanclea lineages. This newly recovered clade may correspond with the genus Zanclella or with a cryptic genus. The lack of both morphological and molecular data for several zancleid species did not allow to address this issue. Additionally, the single-and multi-locus phylogeny reconstructions reveal that both the family Zancleidae and the genus Zanclea are polyphyletic taxa, since they are composed of at least three divergent lineages. Most zancleid species have polyps and medusae similar to other closely related taxa, and this conserved general morphology poses a challenge in the delimitation of species, genera and even families in this group. Consequently, further conjunct morphological and molecular efforts are strongly needed to clarify the diversity and evolution of the family Zancleida as a whole.


Ivan Löbl

The present Catalogue comprises information on 48 genera and 1802 valid species of the coleopterous subfamily Scaphidiinae. Unlike previous catalogues of the group, it gives exact type localities, depositories and sexes of primary types whenever traceable. Genera are listed with their type species and grammatical gender. References are given to subsequent taxonomic and nomenclatural acts, records, and to all other relevant information. Distributional information is provided per country, for larger or insular countries per subunits. An overview of fungus and slime mould hosts is annexed. The references to all taxa have been checked by the author. Infrasubspecific entities and priorities are dealt to comply with the ICZN. Nomenclatural problems are highlighted, and several new records are given. Extinct taxa are listed separately.

Cleopatra M. Loza, Oliver Reutimann, Marcelo R. Sánchez-Villagra, Alfredo A. Carlini and Gabriel Aguirre-Fernández

Edited by A. van der Geer

The systematic value of the middle-ear ossicles, in particular the malleus, has been long recognized for diverse groups. We present systematic work on the characters of the middle-ear ossicles of pinnipeds, focusing on until now poorly studied Southern Hemisphere species. Mallei were extracted from 16 specimens of pinnipeds belonging to five austral and one boreal species of Phocidae and two austral species of Otariidae. Several characters used in this study have been described previously, and some were here modified. Three new characters are here defined and analysed. All characters were mapped onto the phylogeny. Our character analysis shows the transformations that have occurred in the evolution of middle ear ossicles in pinnipeds and identifies diagnostic features of many of its clades. Beyond the identification of specific changes within eachclade, our study of pinniped ossicle evolution documents the occurrence of anatomical convergences with other groups of mammals that live in an aquatic environment, as has occurred in other organ systems as well.

Ondřej Korábek, Lucie Juřičková, Igor Balashov and Adam Petrusek

Edited by T. de Winter

Helix lucorum is a large synanthropic land snail of substantial economic importance, which has been recently reported from a number of new sites in Western, Central, and Eastern Europe. It is an originally Anatolian and Caucasian species, but its presumed natural distribution also covers the south and east of the Balkans. Populations of unclear origin, known as Helix lucorum taurica, live in the south-western part of Crimea. The Balkan and Crimean populations differ in their appearance, were long treated as different species or subspecies, and the Crimean populations are protected by law as a presumably endemic taxon. Here we explore the origins of European populations using phylogenetic analysis of mitochondrial markers. The results point to north-eastern Anatolia and the Lesser Caucasus as the centre of H. lucorum diversification. The Crimean conchological form, along with the associated mitochondrial lineage, is not endemic to that peninsula and was likely introduced there. Other European samples belong to a different lineage, which is associated with the nominotypical conchological form of H. lucorum. The conchological characteristics of the nominotypical form are unusual within the genus Helix, and we propose this reflects a change in habitat use and parallels the evolution of some other lineages of Helicidae. As a result, the typical European H. lucorum differs considerably from the Crimean populations, but the two lineages just represent opposite ends of a continuum in conchological characteristics. Their formal descriptions were based on probably introduced populations detached from the main range. Separated from the geographic pattern of the overall variation of H. lucorum they appeared distinctive, thus confounding the taxonomy of the species. Currently, the division of H. lucorum into subspecies appears unwarranted. Helix lucorum may not be a unique example of a large snail successfully expanding from Anatolia, as we argue that even the native origin of Balkan populations is uncertain. However, further sampling in north-western Turkey and analysis of archaeozoological findings from the south-eastern Balkans is needed to evaluate this hypothesis.

Denis Audo, Matúš Hyžný and Sylvain Charbonnier

Edited by R. Vonk

Polychelidan lobsters, as the sister group of Eureptantia (other lobsters and crabs), have a key-position within decapod crustaceans. Their evolutionary history is still poorly understood, although it has been proposed that their Mesozoic representatives largely inhabited shallow-marine environment and only later sought refuge in deep water. This view has recently been challenged, so the evolutionary history of polychelidans is in a need of re-appraisal. The earliest representatives, such as Tetrachela from the Late Triassic of Austria and Italy, are of great importance because of their potential in investigation of life habits of early polychelidans. Tetrachela lived in a relatively deep water, however, its well-developed eyes suggest an environment where light was still present. With its massive dorsoventrally flattened body plan, Tetrachela was probably benthic; the shape of its mandible and stocky first pereiopods suggest it was a scavenger and/or fed on slowly moving or sedentary animals. The carapace of Tetrachela has a peculiar groove pattern, which leads us to redefine some elements of the nomenclature of grooves used for polychelidans. Based on the present revision we propose that the second incision and its associated groove correspond to the hepatic groove, not the postcervical or the branchiocardiac grooves as interpreted previously. This revision allows us to review the homologies of cephalothoracic groove between polychelidans and other notable groups of decapod crustaceans.

Naturalists in the Field

Collecting, Recording and Preserving the Natural World from the Fifteenth to the Twenty-First Century


Edited by Arthur MacGregor

Interposed between the natural world in all its diversity and the edited form in which we encounter it in literature, imagery and the museum, lie the multiple practices of the naturalists in selecting, recording and preserving the specimens from which our world view is to be reconstituted. The factors that weigh at every stage are here dissected, analysed and set within a historical narrative that spans more than five centuries. During that era, every aspect evolved and changed, as engagement with nature moved from a speculative pursuit heavily influenced by classical scholarship to a systematic science, drawing on advanced theory and technology. Far from being neutrally objective, the process of representing nature is shown as fraught with constraint and compromise.

With a Foreword by Sir David Attenborough

Contributors are: Marie Addyman, Peter Barnard, Paul D. Brinkman, Ian Convery, Peter Davis, Felix Driver, Florike Egmond, Annemarie Jordan Gschwend, Geoff Hancock, Stephen Harris, Hanna Hodacs, Stuart Houston, Dominik Huenniger, Rob Huxley, Charlie Jarvis, Malgosia Nowak-Kemp, Shepard Krech III, Mark Lawley, Arthur Lucas, Marco Masseti, Geoff Moore, Pat Morris, Charles Nelson, Robert Peck, Helen Scales, Han F. Vermeulen, and Glyn Williams.

Valentin Rineau, René Zaragüeta i Bagils and Michel Laurin

Edited by R. Sluys

Simulation-based and experimental studies are crucial to produce factual arguments to solve theoretical and methodological debates in phylogenetics. However, despite the large number of works that tested the relative efficiency of phylogenetic methods with various evolutionary models, the capacity of methods to manage various sources of error and homoplasy has almost never been studied. By applying ordered and unordered methods to datasets with iterative addition of errors in the ordering scheme, we show that unordered coding in parsimony is not a more cautious option. A second debate concerns how to handle reversals, especially when they are regarded as possible synapomorphies. By comparing analyses of reversible and irreversible characters, we show empirically that three-taxon analysis (3ta) manages reversals better than parsimony. For Brownian motion data, we highlight that 3ta is also more efficient than parsimony in managing random errors, which might result from taphonomic problems or any homoplasy generating events that do not follow the dichotomy reversal/ convergence, such as lateral gene transfer. We show parsimony to be more efficient with numerous character states (more than four), and 3ta to be more efficient with binary characters, both methods being equally efficient with four states per character. We finally compare methods using two empirical cases of known evolution.

Roman Croitor and Theodor Obada

Edited by A. van der Geer

This article reports antler remains from the Late Paleolithic site of Climăuți II (Republic of Moldova) confirming the presence of wapiti Cervus canadensis in the Late Pleistocene of Western Eurasia. The occurrence of wapiti in the East Carpathian area by 20 ky BP coincides with the local extinction of Megaloceros giganteus, Crocuta spelaea, and Ursus spelaeus, and substitution of local forest reindeer with grazing tundra-steppe Rangifer tarandus constantini. We here provide an overview of paleontological data and opinions on the presence of Cervus canadensis in Europe, a discussion on the taxonomic status and systematic position of the extinct deer Cervus elaphus palmidactyloceros, and propose a dispersal model for wapiti in Europe during the Late Pleistocene.

D. A. Caraballo and M. S. Rossi

Edited by V. Nijman

Subterranean rodents of the genus Ctenomys have experienced an explosive radiation and rapidly colonized the southern cone of South America. The torquatus group, one of the main groups of the genus, comprises several species and species complexes which inhabit the eastern part of the distribution of Ctenomys including southern Brazil, northern and central Uruguay and north-eastern Argentina. This group has undergone a high chromosomal diversification with diploid numbers varying from 41 to 70. The aim of this study was to investigate the origins of the torquatus group as well as its diversification patterns in relation to geography and cladogenesis. Based on mitochondrial cytochrome b nucleotide sequences we conducted a Bayesian multi-calibrated relaxed clock analysis to estimate the ages of the torquatus group and its main lineages. Using the estimated evolutionary rate we performed a continuous phylogeographic analysis, using a relaxed random walk model to reconstruct the geographic diffusion of the torquatus group in a temporal frame. The torquatus group originated during the early Pleistocene between 1.25 and 2.32 million years from the present in a region that includes the northwest of Uruguay and the southeast of the Brazilian state of Río Grande do Sul. Most lineages have dispersed early towards their present distribution areas going through subsequent range expansions in the last 800,000 – 700,000 years. Ctenomys torquatus went through a rapid range expansion for the last 200,000 years, becoming the most widespread species of the group. The colonization of the Corrientes and Entre Ríos Argentinean provinces supposes at least two crossing events across the Uruguay River between 1.0 and 0.5 million years before the present, in the context of a cold and dry paleoenvironment. The resulting temporal and geographic frame enables the comprehension of the incidence of both, the amplitude of distribution areas and divergence times into the patterns of chromosomal diversification found in the group.

La restauration de la création

Quelle place pour les animaux?


Edited by Michele Cutino, Isabel Iribarren and Françoise Vinel

La restauration de la création se propose d’examiner le statut des animaux dans la pensée chrétienne ancienne et médiévale selon une perspective eschatologique, centrée sur la question du salut des animaux dans le projet divin. L'ouvrage est organisé en trois parties : les sources bibliques, notamment la promesse du renouvellement de la création dans Rm 8, 21 ; les élaborations doctrinales dans la période patristique puis au Moyen Âge ; enfin, des réflexions contemporaines à propos du statut des animaux dans nos sociétés. Souvent absente du débat sociétal à propos des animaux, la perspective historique chrétienne proposée dans ce volume se veut une contribution originale à la réflexion actuelle sur le statut juridique et éthique des animaux.

La restauration de la création aims to examine the status of animals in ancient and medieval Christian thought following an eschatological perspective, that is, focusing on the question of the salvation of animals according to the divine plan. The volume is articulated in three parts: Biblical sources, in particular the promise of the renewal of creation according to Rm 8, 21; Patristic and medieval doctrinal elaborations on the question; finally, contemporary considerations regarding the status of animals in our societies. Often absent from the current social debate on the subject, the historical Christian perspective which this volume proposes is intended as an original contribution to today’s ongoing reflexion on the legal and ethical status of animals.

Editors Contributions to Zoology

Samuel G. Penny, Angelica Crottini, Franco Andreone, Adriana Bellati, Lovasoa M.S. Rakotozafy, Marc W. Holderied, Christoph Schwitzer and Gonçalo M. Rosa

Edited by J.W. Arntzen

Prior herpetological surveys in 1996 and 2000 identified 14 species of amphibians and 32 species of reptiles from the Sahamalaza Peninsula. This work increases the total number of amphibian and reptile species known from this area to 20 and 43 respectively. To maximise our chances of species detection, survey effort covered the entire wet season and part of the dry season, and utilised a combination of opportunistic searching, transect searching, pitfall trapping, and acoustic recording. We identified species through an integrative taxonomic approach, combining morphological, bioacoustic and molecular taxonomy. Together, this enabled the detection of cryptic and seasonally inactive species that were missed in the shorter prior surveys that relied on morphological identification alone. The taxonomic identification of amphibians utilised a fragment of the mitochondrial 16S rRNA gene; taxonomic identification of reptiles utilised a fragment of the mitochondrial COI gene, and when necessary, also mitochondrial fragments of the 16S rRNA ND1, ND2, ND4 genes. All sequences were deposited in Genbank and COI sequences were also deposited in the BOLD database to foster taxonomic identification of malagasy reptiles. We report two new taxa: a species of Boophis, since described as B. ankarafensis, and a candidate new species of microhylid (genus: Stumpffia). We document range expansions of Boophis tsilomaro, Cophyla berara, Blaesodactylus ambonihazo beyond their type localities. Along with significant range expansions across a range of taxa, including Blommersia sp. Ca05, Boophys brachychir, Brookesia minima, Ebenavia inunguis, Geckolepis humbloti, Madascincus stumpffi, Pelomedus subrufa and Phelsuma kochi. Forest in the peninsula is under extreme pressure from human exploitation. Unless unsustainable agricultural and pastoral practices encroaching on these habitats halt immediately, both forest and the species that occur there, several of which appear to be local endemics, may be irreversibly lost.

Kennet Lundin, Tatyana Korshunova, Klas Malmberg and Alexander Martynov

Edited by B.W. Hoeksema

Based on morphological, bathymetric and molecular data comparing recently collected Arctic and North Atlantic specimens with morphological and bathymetrical data on historical museum specimens, a unique relict population of the deep-water mollusc Dendronotus velifer G.O. Sars, 1878 (Gastropoda: Nudibranchia) is shown to have existed in the deepest section of the Swedish Gullmar Fjord (the only true silled fjord in Sweden) at least until the middle of the 20th century. This population is more than 1500 km away from the nearest point in the species’ distributional range in the Arctic Ocean today. Using an integrative approach incorporating the data mentioned above, including genetic distances, from recently collected specimens taken from the Arctic Ocean, D. velifer is validated and its species status is restored, for the first time in more than a century after being regarded as a junior synonym of D. robustus. The bathymetric data for historical and recently collected specimens of D. velifer demonstrate significant differences compared to the shallow-water species D. robustus. The findings support the necessity of a stronger protection for the unique marine habitats of the Gullmar Fjord.

Menno Schilthuizen, Rob Langelaan, Nicola Hemmings, Wesley van Oostenbrugge and Stefan Visser

Edited by A. Minelli

In animals, cell polarity may initiate symmetry breaking very early in development, ultimately leading to whole-body asymmetry. Helical sperm cells, which occur in a variety of animal clades, are one class of cells that show clearly visible bilateral asymmetry. We used scanning-electron microscopy to study coiling direction in helical sperm cells in two groups of animals that have figured prominently in the sperm morphology literature, namely land snails, Stylommatophora (514 spermatozoa, from 27 individuals, belonging to 8 species and 4 families) and songbirds, Passeriformes (486 spermatozoa, from 26 individuals, belonging to 18 species and 8 families). We found that the snail sperm cells were consistently dextral (clockwise), whereas the bird sperm cells were consistently sinistral (counterclockwise). We discuss reasons why this apparent evolutionary conservatism of sperm cell chirality may or may not be related to whole-body asymmetry.

Jacques J. M. van Alphen and Jan W. Arntzen

Edited by Alessandro Minelli

In a recent paper it was suggested that results published by Kammerer (1911) on the midwife toad could be explained by epigenetics (Vargas et al., 2016). We show that data thought to be fitting are based on untested assumptions about the underlying genetic mechanisms. We cite recent studies on the genetics of life history traits, in particular egg-size and number, to show that these assumptions are not realistic. We review aspects of Kammerer’s experimental results on the midwife toad for which there are no plausible mechanisms, i.e. toads switching from land-breeding to water-breeding in response to an increase in temperature, eggs becoming resistant to moulds within a few generations, the gradual development of nuptial pads, heterochronous changes in the development of water-born larvae, and conclude that Kammerer cannot have obtained the results he claims. We argue that natural selection would not have favoured a change in reproductive mode and the loss of parental care and that an epigenetic master switch, affecting many different traits simultaneously, would have either eroded during more than 15 million years of land-breeding and/or would have disappeared by natural selection against it. Finally, we show that Kammerer's data are remarkably close to the invoked Mendelian ratio and too good to be genuine. We conclude that Kammerer’s data are fictitious and that Vargas et al. (2016) have used non-existing data in search of support for a role of epigenetics in neo-Lamarckian evolution.

Daniel Martin, Miguel A. Meca, João Gil, Pilar Drake and Arne Nygren

Edited by B.W. Hoeksema

Oxydromus humesi is an annelid polychaete living as a strict bivalve endosymbiont (likely parasitic) of Tellina nymphalis in Congolese mangrove swamps and of Scrobicularia plana and Macomopsis pellucida in Iberian saltmarshes. The Congolese and Iberian polychaete populations were previously considered as belonging to the same species, the latter showing regular distribution, intra-specific aggressive behaviour, and complex hostentering behaviour. The fresh Iberian samples enabled us to undertake consistent morphometric analyses, as well as to further analyse the characteristics of the association and the population dynamics of the Iberian population hosted by S. plana. Among the morphological differences between the Congolese and Iberian specimens, leading to the description of the latter as Oxydromus okupa sp. nov., the most important are: 1) longer cephalic appendages, 2) greater distance between the eyes, 3) larger dorsal cirrostyle in relation to the corresponding dorsal lobe and cirrophore. Moreover, dorsal and ventral lobes are similar in length, with the tip of the former reaching the tip of the latter in O. okupa sp. nov., while the dorsal lobe is much shorter than the ventral one in O. humesi. Mature adults of O. okupa sp. nov. occurred during the whole study period, with a higher percentage of ripe females in spring and, particularly, in summer. Numerous host specimens showed the symbiont’s most preferred shell length (>26 - 36 mm). However, the prevalence was very low (usually <5%) and showed a clear seasonal pattern, being lower during spring/summer. This suggests that males are able to leave their hosts during this period, most likely to improve fertilization by directly entering or approaching a host occupied by a ripe female, while females usually remain inside. Based on the new results, the current knowledge of symbiotic Hesionidae and their relationships with invertebrate hosts is updated and discussed.

José A. Jurado-Rivera, Genaro Álvarez, José A. Caro, Carlos Juan, Joan Pons and Damià Jaume

Edited by R. Vonk

The molecular systematics of the subterranean amphipod genus Haploginglymus is addressed through the phylogenetic analysis of three DNA gene fragments (nuclear ribosomal 28S and protein- coding Histone 3, plus mitochondrial Cytochrome c Oxidase subunit I). We take advantage of the description of a new species from southern Spain (Haploginglymus geos sp. nov.) to assess the singularity of this genus endemic to the Iberian Peninsula and the inclusion of the morphologically aberrant H. morenoi within Haploginglymus. Our results corroborate the monophyly of the family Niphargidae but shows Niphargus to be paraphyletic as it currently stands, with Haploginglymus appearing nested within it. A strongly supported sister-group relationship between niphargids and the (thalassoid) pseudoniphargids is recovered as well, but we propose the Niphargidae should continue to be considered as a primary limnic group for biogeographic purposes despite its presumed marine derivation. Our findings are in agreement with previous studies that suggest the family Niphargidae originated in the late Cretaceous in the NE Atlantic, from where it eventually expanded across continental Europe.

Christina Nagler, Jens T. Høeg, Carolin Haug and Joachim T. Haug

Edited by R. Vonk

The larval phase of metazoans can be interpreted as a discrete post-embryonic period. Larvae have been usually considered to be small, yet some metazoans possess unusually large larvae, or giant larvae. Here, we report a possible case of such a giant larva from the Upper Jurassic Solnhofen Lithographic limestones (150 million years old, southern Germany), most likely representing an immature cirripede crustacean (barnacles and their relatives). The single specimen was documented with up-to-date imaging methods (macro-photography, stereo-photography, fluorescence photography, composite imaging) and compared with modern cirripede larvae. The identification is based on two conspicuous spine-like extensions in the anterior region of the specimen strongly resembling the so-called fronto-lateral horns, structures exclusively known from cirripede nauplius larvae. Notably, at 5 mm in length the specimen is unusually large for a cirripede nauplius. We therefore consider it to be a giant larva and discuss possible ecological and physiological mechanisms leading to the appearance of giant larvae in other lineages. Further findings of fossil larvae and especially nauplii might give new insights into larval evolution and plankton composition in the past.

Xin Tong, Lu Jiang and Bao-Zhen Hua

Edited by Herman de Jong

Sexually reproductive insects exhibit diverse mating behaviors. However, the mating pattern remains unknown for Panorpodes of Panorpodidae to date. In this study, we investigated the mating behavior and copulatory mechanism of the short-faced scorpionfly Panorpodes kuandianensis Zhong, Zhang and Hua, 2011 for the first time. The results show that the male provides a salivary mass as a nuptial gift to the female and starts to copulate with the female in a V-shaped position, then changes to an end-to-end position by temporarily twisting the female abdominal segments VII−IX by 180°. During mating the basal processes and the basal teeth of the gonostyli and the hypandrium are used to obtain copulation and sustain the coupling of genitalia to secure successful sperm transfer. This unique mating pattern is greatly different from that of other Mecoptera reported and is likely evolved as an adaptation in the context of sexual conflict.

Marta Guntiñas, Jorge Lozano, Rodrigo Cisneros, Carlos Narváez and Jorge Armijos

Edited by V. Nijman

This study details for the first time the food habits of the culpeo or Andean fox (Lycalopex culpaeus) in high areas of the Ecuadorian Andes during a complete annual cycle. Our objective was to identify prey species, calculate their contribution to diet and test for temporal variation. In total, 304 Andean fox scats were collected within the Podocarpus National Park (Loja province, southern Ecuador). By analysing the content of scats 413 prey items were identified and classified into eight prey groups. Sub-sequently, the frequency of occurrence (FO) for each prey group was calculated per month together with the relative contribution of estimated consumed biomass (CB). In addition, the Shannon-Wiener’s index was calculated as a measurement of monthly diet diversity. The results show a diet dominated by cervids belonging to the Mazama and Pudu genus (70% FO), followed by small mammals (30% FO), large rodents (12% FO), carnivorous species (10% FO) and lagomorphs (8% FO). There was temporal variation in the consumption of both deer and small mammals. Cervids also provided the major part of the consumed biomass (70% CB), the remaining prey categories each accounted for less than 11% CB. Negative correlations in consumption were observed between cervids and two other groups, rabbits and small mammals. Trophic diversity values fluctuated throughout the year showing a mean of 1.7. The results obtained show a diets very different from those previously reported; deer being the bulk of the diet instead of rodents or other mammals. The consumption of carnivorous species in the area was also high in comparison with other regions. Overall, this study clearly demonstrates that the culpeo is an important top predator in the high-Andean ecosystem.

Nikolai Y. Neretin, Anna E. Zhadan and Alexander B. Tzetlin

Edited by R. Vonk

In the present study, we investigated the biology of Dyopedos bispinis, a mast-building amphipod that is abundant near the N. Pertsov White Sea Biological Station. To examine the peculiarities of mast building in Dyopedos bispinis, we studied the social structure of individuals inhabiting the masts and identified the preferred substrata through underwater photography and direct observations, characterized the internal and external structures of the masts, and studied the ultrastructure of pereopodal silk glands using scanning and transmission electron microscopy (SEM and TEM, respectively). The most frequent substrata for mast building are other fouling organisms, including hydroids, bryozoans, ascidians and sponges. As in other corophiids, each Dyopedos bispinis mast represents the territory of one female and, occasionally, one male, but unique collective masts occupied by three or more (up to 23) adults were also observed. Masts comprise one or 2-4 central cylinders and a laminated cortex that contains detritus and amphipod silk layers. The pereopodal glandular complex of Dyopedos bispinis is composed of two distinct gland groups, proximal and distal, in each pereopod 3-4, and ducts in the glandular complex lead into a common chamber in the dactylus. The proximal glands are multicellular; their secretory cells are uninuclear, unlike in certain other amphipods; and the cell membrane is deeply invaginated. The invaginations are filled with extensions of the cytoplasm of lining cells, but the origin of the lining is unclear. Axon terminals were observed adjacent to the secretory cells, and it is assumed that these axons regulate amphipod silk glands. The proximal silk glands of Dyopedos bispinis have similarities with the lobed and rosette glands of isopods, but they have strongly elongated forms. We refer to these glands as pseudotubular glands. Such glands are rarely observed in Crustacea and have only been described in silk-producing pereopodal systems of marine Peracarida and in the antenna of terrestrial Malacostraca.


Wilfried R. Arnscheid and Michael Weidlich

In Psychidae Arnscheid and Weidlich provide for the first time a complete tool for identifying the European bag worm moths. The book will provide a sufficient overview of the systematics and distribution of the European Psychidae. A total of 246 species is recognized. Description and diagnoses are accompanied by colour figures of the adults, usually depicting variation of male and female if the latter are winged. Black and white photographs of the male genitalia of most species (excluding parts of Naryciinae and Taleporiinae due to their similarity) are given for the first time. Notes on distribution and bionomics are added for every species. One new subfamily, one new genus and three new species are described.


Wilfried R. Arnscheid and Michael Weidlich


Wilfried R. Arnscheid and Michael Weidlich


Wilfried R. Arnscheid and Michael Weidlich

Gerrit Potkamp, Mark J.A. Vermeij and Bert W. Hoeksema

Edited by Danwei Huang

Snails of the genus Coralliophila (Muricidae: Coralliophilinae) are common corallivores in the Caribbean, feeding on a wide range of host species. In the present study, the morphological and genetic variation in C. galea and C. caribaea were studied in relation to their association with host coral species at Curaçao. Differences in shell shape among snails living on different hosts were quantified using geometric morphometric and phylogenetic relationships were studied using two mitochondrial markers (12S and COI). Based on these analyses, a new species, C. curacaoensis sp. nov., was found in association with the scleractinian coral Madracis auretenra. Both C. galea and C. caribaea showed host-specific differences in shell shape, size, and shell allometry (i.e. changes in morphological development during growth). Shell spire variability contributed foremost to the overall variation in shell shape. In C. caribaea minor genetic differences existed between snails associated with scleractinian and alcyonacean corals, whereas in C. galea such intraspecific variation was not found. These results shed more light on morphological and genetic differences among coral-associated fauna living on different host species.


Wilfried R. Arnscheid and Michael Weidlich


Wilfried R. Arnscheid and Michael Weidlich


Wilfried R. Arnscheid and Michael Weidlich


Wilfried R. Arnscheid and Michael Weidlich


Wilfried R. Arnscheid and Michael Weidlich


Wilfried R. Arnscheid and Michael Weidlich

Roberto Guidetti, Sandra J. McInnes, Michele Cesari, Lorena Rebecchi and Omar Rota-Stabelli

Edited by Alessandro Minelli

The origin of the Antarctic continental extant fauna is a highly debated topic, complicated by the paucity of organisms for which we have clear biogeographic distributions and understanding of their evolutionary timescale. To shed new light on this topic, we coupled molecular clock analyses with biogeographic studies on the heterotardigrade genus Mopsechiniscus. This taxon includes species with endemic distributions in Antarctica and other regions of the southern hemisphere. Molecular dating using different models and calibration priors retrieved similar divergence time for the split between the Antarctic and South American Mopsechiniscus lineages (32–48 Mya) and the estimated age of the Drake Passage opening that led to the separation of Antarctica and South America. Our divergence estimates are congruent with other independent studies in dating Gondwanan geological events. Although different analyses retrieved similar results for the internal relationships within the Heterotardigrada, our results indicated that the molecular dating of tardigrades using genes coding for ribosomal RNA (18S and 28S rDNA) is a complex task, revealed by a very wide range of posterior density and a relative difficulty in discriminating between competing models. Overall, our study indicates that Mopsechiniscus is an ancient genus with a clear Gondwanan distribution, in which speciation was probably directed by a cooccurrence of vicariance and glacial events.

Isabel T. Hyman, Irantzu de la Iglesia Lamborena and Frank Köhler

Edited by S.E.T. van der Meij

The south-eastern Australian helicarionid clade currently comprises six genera of snails and semislugs united by genital characters, including an epiphallic flagellum that produces a spiraling, spinose spermatophore, the absence of an epiphallic caecum, and the presence of at most a very short vagina. We comprehensively revise the taxonomy of this group based on comparative analyses of key morphological features and mitochondrial markers COI and 16S, revise the placement of several species described recently on the basis of shell morphology alone, and describe new taxa. The snail genus Brevisentis is monophyletic as currently understood, but includes an additional undescribed species from Wollemi National Park, New South Wales. Mysticarion is shown to contain four arboreal semislugs with wide, disjunct ranges, including one new species (Mysticarion obscurior sp. nov.). We synonymise Fastosarion staffordorum with Mysticarion hyalinus. The semislug Desidarion is synonymised with Parmavitrina, and we describe two new species of this genus, P. flavocarinata sp. nov. and P. maculosa sp. nov. The semi-arboreal semislug Cucullarion is herein included in the south-eastern Australian helicarionid clade based on morphological and genetic evidence, despite its more northerly distribution. Two small semislugs so far placed in Peloparion do not group together and a new genus, Ubiquitarion gen. nov., is described for Peloparion iridis. All of these genera, together with the semislug Helicarion (not included here), form a monophyletic radiation endemic to southeastern Australia.


A Compendium of the Geographical Distribution and a Summary of Their Biology


Lourelle Neethling and Annemariè Avenant-Oldewage

This is a compendium of current knowledge about the crustacean subclass Branchiura Thorell, 1864. An overview of the group is presented, starting from the first species description, and reports of taxonomic changes. It also provides a condensed retrospect of each genus and includes the characteristics of each genus, the geographical distribution of each species arranged according to occurrence per continent; and aspects of the anatomy, physiology, host-parasite interactions and phylogeny are discussed. In order to condense the information available on members of the subclass, additional literature sources on each aspect are tabulated. This text will be useful for fish health practitioners, researchers and students of Parasitology and Fish Veterinary Medicine.

The contents of this volume were originally published in 2016 in Crustaceana volume 89, issue 11-12.

Vincent Nijman and Daniel Bergin

Edited by S.E.T. van der Meij

Reptiles are traded globally for medicinal purposes. Historic qualitative accounts of reptiles used as medicine in Morocco are numerous, but contemporary quantitative data are rare. In 2013- 2014, we surveyed 49 wildlife markets in 20 towns throughout Morocco, plus the Spanish exclaves of Ceuta and Melilla. We recorded 1,586 specimens of at least nine species for sale in 14 of the Moroccan markets with a combined value of about US $100,000. The most prominent markets were those in Marrakesh, Meknes, Casablanca, and Fez, with the former two cites trading equal quantities of dried and live specimens and the latter two trading mainly dried specimens. Common species were the Mediterranean chameleon (Chamaeleo chamaeleon) with 720 specimens (506 dried, 214 alive) and the Bell’s Dabb lizard (Uromastyx acanthinura) with 428 specimens (247 dried, 181 alive), both traded in 10 markets, and spur-thighed tortoise (Testudo graeca; 57 carapaces), for sale in eight cities. Over 200 African rock python (Python sebae) skins were identified and may have been illicitly imported from other parts of Africa. The turnover of Mediterranean chameleon and Bell’s Dabb lizard specimens after four weeks as measured by repeat surveys was 66% for both species, resulting in an estimated annual turnover of 1,520 chameleons (range 921–2,303) and 775 lizards (range 364–1,174). Despite legal protection and regulations locally within Morocco and internationally through CITES, reptiles are commonly and openly traded for medicinal purposes throughout Morocco. However, traders are not forthcoming in conveying the legal status of these species and restrictions on trade to potential buyers. Increased enforcement of existing wildlife protection legislation is needed to prevent this exploitation from the illegal wildlife trade that could negatively impacts imperiled species.

Tania Trujillo, Jorge Gutiérrez-Rodríguez, Jan W. Arntzen and Iñigo Martínez-Solano

Edited by A. Ivanović

The use of hyper-variable markers across species is often hindered by low cross-species amplification success, a reduced level of polymorphism or a high frequency of null alleles. However, optimizing sets of reliable and informative markers that can be consistently amplified and scored across taxa is key to address questions about patterns of genetic diversity and structure, hybridization and speciation. Here we present 14 newly developed microsatellite markers in the Spined toad (Bufo spinosus), assess their polymorphism in two Iberian populations and test for cross-species amplification in the closely related Common toad (Bufo bufo). We then use the 12 loci co-amplifying in both species to the study of a morphologically intermediate population (Moyaux) from the contact zone in northwest France as well as reference populations of the two species from both sides of the contact zone. Individuals from Moyaux had mtDNA haplotypes of the two species and were identified as hybrids in analyses with software NewHybrids. These results provide solid evidence for ongoing hybridization between B. bufo and B. spinosus, with no apparent restrictions to gene flow.

R.G. Bina Perl, Sarig Gafny, Yoram Malka, Sharon Renan, Douglas C. Woodhams, Louise Rollins-Smith, James D. Pask, Molly C. Bletz, Eli Geffen and Miguel Vences

Edited by J.W. Arntzen

Dramatic global amphibian declines have recently led to an increased concern for many species of this animal class. The enigmatic Hula painted frog (Latonia nigriventer), the first amphibian to be declared extinct but unexpectedly rediscovered in 2011, has remained one of the rarest and most poorly understood amphibians worldwide. Gathering basic biological information on this species, along with an understanding of its disease-related threats remains fundamental for developing risk assessments and conservation strategies. Our surveys in recent years confirmed that L. nigriventer is a localised species with elusive habits. The species appears to follow an opportunistic breeding phenology and has a tadpole morphology similar to its well-studied sister group Discoglossus. However, the adults’ extended annual presence in the aquatic habitat is a major difference from species of Discoglossus. We detected the amphibian chytrid fungus, Batrachochytrium dendrobatidis (Bd), in northern Israel and on Hula painted frogs but did not observe any signs of chytridiomycosis in this species. Our preliminary data on aspects of the innate immunity of L. nigriventer suggest that the skin mucosome of this species contains antimicrobial peptides and a bacterial community differing from other syntopic frogs (Pelophylax bedriagae). The combined knowledge of both natural history and innate immunity of L. nigriventer provides valuable insights to direct future research and conservation management of this critically endangered frog species.

Bregje W. Brinkmann and Charles H.J.M. Fransen

Edited by R. Vonk

Periclimenes rathbunae Schmitt, 1924 is a western Atlantic symbiotic shrimp species mainly associating with anemones. Adult shrimps of P . rathbunae are characterised by an orange-white spotted colour pattern. During fieldwork along the coast of Curaçao (2013), morphologically similar, though generally smaller sized shrimps were collected from the stony coral Dendrogyra cylindrus Ehrenberg, 1834. These specimens were overall more translucent. This study elucidates the taxonomic status of the coral-associated specimens in relation to anemone-associated specimens of P. rathbunae and related anemone-associated species using one nuclear (histone H3, H3), a mitochondrial protein-coding (cytochrome oxidase subunit I, COI) and a mitochondrial ribosomal RNA (16S) gene. Molecular phylogenetic reconstructions clustered the coral-associated specimens with anemone-associated P. rathbunae within a distinct clade, revealing that the aberrant coral-associated shrimp specimens belong to P. rathbunae. It can be concluded that 1) the stony coral D. cylindrus is a host of P. rathbunae, constituting the first record of an association between a scleractinian coral and a palaemonid shrimp species in the Atlantic Ocean; 2) the colour pattern of P. rathbunae is a phenotypically plastic characteristic that varies with size and is depending on the host species; and 3) mean body size of P. rathbunae is smaller on D. cylindrus than on the anemone Stichodactyla helianthus. This raises interesting questions about patterns of host use for this species and warrants further in-depth field ecological study for this species.

Jacques J.M. van Alphen and Jan W. Arntzen

Edited by J. van Rooijen

The Austrian biologist Paul Kammerer (1880-1926) would by now be long forgotten if Arthur Koestler had not published ‘The case of the Midwife toad’, in which he depicted Kammerer as a victim of the paradigm battle between neo-Darwinists and Lamarckists. Kammerer is still on the scientific agenda, with at least 10 publications since 2005. The question is still out if Kammerer fabricated his scientific results or not. In this paper we provide the evidence that Kammerer consistently faked experimental results. We show (1) that the design of his experiments could never have produced the results that he claimed, (2) that the assumptions he made about the developmental biology of the species he studied are falsified by recent research, and (3) that the specimens he showed as proof for the success of his experiments came from nature.

Deborah Wall-Palmer, Alice K. Burridge, Katja T.C.A. Peijnenburg, Arie Janssen, Erica Goetze, Richard Kirby, Malcolm B. Hart and Christopher W. Smart

Edited by D. Huang

The genus Protatlanta is thought to be monotypic and is part of the Atlantidae, a family of shelled heteropods. These microscopic planktonic gastropods are poorly known, although research on their ecology is now increasing in response to concerns about the effects of ocean acidification on calcareous plankton. A correctly implemented taxonomy of the Atlantidae is fundamental to this progressing field of research and it requires much attention, particularly using integrated molecular and morphological techniques. Here we use DNA barcoding, shell morphology and biogeography to show that the genus Protatlanta includes at least two valid species in the Atlantic Ocean. Protatlanta souleyeti and Protatlanta sculpta were found to be separate species, with different shell morphology and separated by a K2P genetic distance of 19% sequence divergence at the Cytochrome Oxidase 1 gene. This evidence supports the revival of the species name P. sculpta, which was described by Issel in 1911, but has not been recognised as a valid species since 1915


Edited by Annie Potts

The analysis of meat and its place in Western culture has been central to Human-Animal Studies as a field. It is even more urgent now as global meat and dairy production are projected to rise dramatically by 2050. While the term ‘carnism’ denotes the invisible belief system (or ideology) that naturalizes and normalizes meat consumption, in this volume we focus on ‘meat culture’, which refers to all the tangible and practical forms through which carnist ideology is expressed and lived. Featuring new work from leading Australasian, European and North American scholars, Meat Culture, edited by Annie Potts, interrogates the representations and discourses, practices and behaviours, diets and tastes that generate shared beliefs about, perspectives on and experiences of meat in the 21st century.

Roberto Arrigoni, Francesca Benzoni, Danwei Huang, Hironobu Fukami, Chaolun Allen Chen, Michael L. Berumen, Mia Hoogenboom, Damian P. Thomson, Bert W. Hoeksema, Ann F. Budd, Yuna Zayasu, Tullia I. Terraneo, Yuko F. Kitano and Andrew H. Baird

Edited by R.W.M. van Soest

The scleractinian family Lobophylliidae is undergoing a major taxonomic revision thanks to the combination of molecular and morphological data. In this study, we investigate the evolutionary relationships and the macro- and micromorphology of six nominal coral species belonging to two of the nine molecular clades of the Lobophylliidae, clades A and B, and of Symphyllia wilsoni, a lobophylliid species analyzed from a molecular point of view for the first time. Sequence data from mitochondrial DNA (COI and the intergenic spacer between COI and l-rRNA), and nuclear DNA (histone H3 and ITS region) are used to generate robust molecular phylogenies and a median-joining haplotype network. Molecular results are strongly in agreement with detailed observations of gross- and fine-scale morphology of skeletons, leading to the formal revision of the genera Micromussa and Homophyllia and the description of two newly discovered zooxanthellate shallow-water species, Micromussa pacifica sp. nov. Benzoni & Arrigoni and Micromussa indiana sp. nov. Benzoni & Arrigoni, and a new genus, Australophyllia gen. nov. Benzoni & Arrigoni. In particular, Acanthastrea lordhowensis and Montastraea multipunctata are moved into Micromussa, A. hillae is synonymized with A. bowerbanki and is transferred to Homophyllia, and a revised diagnosis for both genera is provided. Micromussa pacifica sp. nov. is described from the Gambier Islands with its distribution spanning New Caledonia and eastern Australia. Despite a superficial resemblance with Homophyllia australis, it has distinctive macroand micromorphological septal features. Micromussa indiana sp. nov., previously identified as M. amakusensis, is here described from the Gulf of Aden and the southern Red Sea as a distinct species that is genetically separated from M. amakusensis and is morphologically distinct from the latter due to its smaller corallite size and lower number of septa. Finally, molecular trees show that S. wilsoni is closely related, but molecularly separated from clades A and B, and, also based on a unique combination of corallite and sub-corallite characters, the species is moved into Australophyllia gen. nov. These findings confirm the need for using both genetic and morphological datasets for the ongoing taxonomic revision of scleractinian corals.

Jonas Keiler, Stefan Richter and Christian S. Wirkner

Edited by R. Vonk

The Southern Australian crustacean species Lomis hirta (Lomisoidea: Lomisidae) is a representative of one of the three anomuran taxa which obtained their crab-like habitus independently from each other. This process, the evolutionary transformation into a crab-like form, is termed carcinization. To shed light on the morphological changes which took place during carcinization and to investigate structural dependence (coherence) between external and internal morphological characters, we studied L. hirta and representatives of its putatively most closely related taxa, Aegla cholchol (Aegloidea: Aeglidae) and Kiwa puravida (Chirostyloidea: Kiwaidae). External and internal anatomy was studied using microcomputertomography and computer-aided 3D reconstruction. A. cholchol and K. puravida belong to equally exceptional lineages: Aegla is endemic to South America and lives in freshwater habitats; Kiwa is a deep sea dweller associated to chemosynthetic bacteria found in methane seeps or hydrothermal vents. On the basis of recent cladistic analyses we reconstruct the anatomical ground pattern of the squat lobster-like last common ancestor of the three taxa and trace the morphological transformations that affected inner and outer morphology in the recent forms. Our results show, among other things, that the pleon in Lomis underwent drastic modifications in the context of carcinization, including a reduction of the muscular portion leaving more room for the hepatopancreas and gonads, and a narrowing of the pleonal ganglia which became shifted anteriorly into the cephalothorax and attached to the cephalothoracic ganglion. We interpret these anatomical changes in Lomis to have come about because of the loss of the caridoid escape reaction, which in turn was a direct consequence of the evolution of a strongly bent pleon as part of the crab-like habitus, and of a hidden lifestyle.

Iva Njunjić, Michel Perreau, Kasper Hendriks, Menno Schilthuizen and Louis Deharveng

Edited by J.C. Biesmeijer

The subtribe Anthroherponina form an iconic group of obligate cave beetles, typical representatives of the Dinaric subterranean fauna, which is considered to be the richest in the world. Phylogenetic studies within this subtribe are scarce and based only on morphological characters, which, due to troglomorphic convergence, are frequently unreliable. Moreover, morphological stasis and morphological polymorphism make classification of taxa difficult. To test if characters that have traditionally been accepted as informative for Anthroherponina classification are indeed reliable, we evaluated the monophyly of the most speciesrich genus of this subtribe - Anthroherpon Reitter, 1889. Our study, based on a molecular phylogenetic analysis of fragments of the 18S, 28S, and COI (both 5’ and 3’ end) loci revealed that the genus Anthroherpon as conventionally defined is polyphyletic. To resolve this polyphyly, we defined one new additional genus, Graciliella n. gen., for which we then examined the intrageneric diversity using molecular and morphometric approaches. Molecular phylogenetic analysis of two COI mitochondrial gene fragments revealed the presence of four species inside Graciliella n. gen., including two new species, which we here describe as G. kosovaci n. sp. and G. ozimeci n. sp. To analyze interspecific morphological differences within Graciliella we performed a discriminant analysis based on 40 linear morphometric measurements. The results showed that differences between species and subspecies inside Graciliella, however subtle they may seem, are measurable and reproducible. All species of the genus are briefly diagnosed, an identification key is proposed and a distribution map of all taxa of Graciliella is provided.

Rob W. M. van Soest

Edited by L.P. van Ofwegen

Sponges from the Kara Sea (Arctic Russia) collected by the Dutch Polar Expedition 1882-83 are identified and discussed. The expedition experienced an unfortunate course of events, as the research vessel RV ’Varna’ was ice-bound and eventually got crushed by ice, leaving the crew and the scientists stranded on an ice floe drifting around in the Kara Sea. The scientists still managed to do many observations and made collections of bottom animals lowering collecting gear through a hole in the ice. After being enclosed for more than a year in the ice, the men travelled in small boats and sledges to the mainland of Russia and from there returned home. The zoological collections were brought to Copenhagen by the Danish research vessel RV ‘Dijmphna’, from where the material was picked up by the Dutch scientists. Most animal groups were subsequently discussed in a series of papers published in the Bijdragen tot de Dierkunde volume 14 (1887). The sponges of the Dutch Polar Expedition were in a rather bad condition when they arrived in Amsterdam and possibly for that reason were largely ignored. They nevertheless appear to be interesting and are here presented as a belated contribution to the 1887 issue, including the description of a species new to science, Lycopodina ruijsi sp. nov. The results are discussed in the framework of our present knowledge of the sponge fauna of the Kara Sea. In Appendix ten already known species are briefly described and SEM images of their spicules provided as a contribution to Kara Sea sponge morphology,

Michelangelo S. Moerland, Chad M. Scott and Bert W. Hoeksema

Edited by D. Huang

Corallivorous Drupella (Muricidae) snails at Koh Tao are reported to have extended their range of prey species following a major coral bleaching event in 2010. Populations of their preferred Acropora prey had locally diminished in both size and abundance, and the snails had introduced free-living mushroom corals in their diet. Although the coral community largely recovered, the Drupella population grew and reached outbreak proportions. For this study, corallivorous muricids at Koh Tao were studied more intensively to examine their identities, distribution and prey choice four years after the bleaching event. Drupella rugosa was identified as the major outbreak species and occurred at densities > 3 m-2 in depth ranges of 2-5 and 5-8 m. The density of D. rugosa was related to the live coral cover, Acropora colony density, and depth. Resource selection ratios revealed that species of Acropora, Psammocora and Pavona corals were attacked more frequently than would be expected based on their availability. Strikingly, fungiid corals were now avoided as prey in the recovered coral community, despite them being part of the preferred diet directly after the bleaching. Although D. rugosa showed a clear prey preference, it appears to be plastic by changing with prey availability. The muricids Drupella margariticola and Morula spinosa occurred in much lower densities and were less often associated with corals. Snails of the opportunistic corallivore D. margariticola usually co-occurred in D. rugosa aggregations, although they also formed feeding aggregations by themselves. Whether M. spinosa generally associates with corals as a corallivore or a scavenger has yet to be determined. Molecular analyses did not reveal cryptic speciation among snails sampled from different coral hosts and also no geographic variation. The present study also showed that corallivory is more common among D. margariticola and M. spinosa than previously known.

Camila Falcione, Alejandra Hernando, Diego Andrés Barrasso and Diego Omar Di Pietro

Edited by J. van Rooijen

The karyotypes of four Xenodontini snake species, Lygophis dilepis, L. meridionalis, L. flavifrenatus and L. anomalus, are here described for the first time. We studied specimens from northeastern Argentina using conventional and silver (Ag-NOR) staining. While the typical ophidian karyotype is 2n = 36, we found that the karyotype of the studied species is 2n = 34, with metacentric and submetacentric chromosome pairs. The 4th NOR staining revealed that nucleolar organizer regions (NORs) are located on one pair of microchromosomes. In L. dilepis and L. anomalus the 4th chromosome pair is heteromorphic, and we suggest that it might be considered as the ZW sex chromosome pair. The optimization of available karyological data on a molecular phylogenetic tree of the tribe Xenodontini shows that the diploid numbers 2n = 28, 30 and 34 represent putative synapomorphy for Erythrolamprus, Xenodon and Lygophis, respectively. Our results provide new insights which fill gaps in our knowledge on the cytology in the genus Lygophis and identified a possible diagnostic character for the genus.

Tina Klenovšek and Vida Jojić

Edited by A. Ivanović

We explored modularity and morphological integration of the ventral cranium during postnatal ontogeny in Martino’s vole (Dinaromys bogdanovi). Two closely related phylogenetic groups, originating from the Central and Southeastern part of the species range in the western Balkans, were considered. As expected, both phylogroups had similar patterns of ontogenetic changes in cranial size and shape variation, modularity and integration. At the level of within individual variation, the hypothesis that the viscerocranial and neurocranial regions are separate modules was rejected, indicating that the hypothesized modules are not developmental, but rather functional. At the level of among individual variation, the viscerocranium and the neurocranium could not be recognized as separate modules at the juvenile stage. The strength of association between the hypothesized modules becomes lower with age which finally results in a clear 2-module organization of the ventral cranium at the adult stage. On the other hand, patterns of morphological integration for the cranium as a whole, the viscerocranium and the neurocranium stay consistent across ontogenetic stages. The developmental mechanism producing integration of the cranium as a whole, as well as integration of the neurocranium, varies throughout postnatal ontogeny. In contrast, we detected the ontogenetic stability of the mechanism responsible for covariation of viscerocranial traits which could provide ongoing flexibility of the viscerocranial covariance structure for high functional demands during lifetime. Findings from our study most likely support the idea of the ‘palimpsest-like’ model of covariance structure. Moreover, similarity or dissimilarity in the patterns of within and among individual variation in different sets of analyzed traits and comparisons across ontogenetic stages demonstrate how studies on small mammals other than mice can give new insights into postnatal cranial development.

Bert Van Bocxlaer and Ellen E. Strong

Edited by D. Huang

The anatomy, functional morphology and evolutionary ecology of the Viviparidae, and the subfamily Bellamyinae in particular, are incompletely known. Partly as a result, genealogical relationships within the family remain poorly understood. Because of this lack in knowledge, few informed hypotheses exist on ancestral states, how differences in body plans between the subfamilies evolved, and how the peculiar biogeographic distribution patterns of viviparids have arisen. Here we document the anatomy, morphology, life history and systematics of Cipangopaludina japonica, a Japanese species that has been introduced into North America, to resolve taxonomic confusion and to improve our understanding of how form and function are related in bellamyines. Anatomical and histological examinations demonstrate marked differences between C. japonicaand other bellamyines in the radula, salivary gland, kidney, nerve ring and reproductive organs. Substantial differences also exist between male and female body organization, but conchological differences between sexes in semi-landmark morphometric analyses are limited. The volume of the brood pouch of females, and hence body and shell size, appear to be good predictors of reproductive success, and the species’ ecological versatility may relate to high fecundity and the ability to alternate between feeding modes. Comparing our observations on C. japonicawith other viviparids and basal Architaenioglossa, we identify several persistent misinterpretations in the literature on how form and function are related in viviparids, not in the least as to female reproductive anatomy. Our reinterpretations improve understanding of the evolution of Viviparidae and its subfamilies, and hopefully will allow future workers to isolate key traits that shaped the evolution of viviparids at the taxonomic levels of their interest for more detailed studies.

Eduardo Ascarrunz, Jean-Claude Rage, Pierre Legreneur and Michel Laurin

Edited by R. Vonk

Triadobatrachus massinotiis a batrachian known from a single fossil from the Early Triassic of Madagascar that presents a combination of apomorphic salientian and plesiomorphic batrachian characters. Herein we offer a revised description of the specimen based on X-ray micro-tomography data. We report previously unknown caudal vertebrae, possible mentomeckelians, and hidden parts of other structures. We also confirm the presence of a ventrolateral ledge on the opisthotic, and we rectify some previous interpretations. There are no cervical ribs and the jaw may have had an angular. The presacral region is composed of 15 vertebrae with a unique atlas-axis complex instead of 14 vertebrae with a bipartite atlas. The configuration of the pelvic girdle is not very clear, although it is likely more plesiomorphic than the anuran- like condition previously assumed. Our re-assessment of the saltatorial performance of Triadobatrachussupports the traditional interpretation that this animal was not a specialised jumper. In order to assess the sequence of events in the early evolution of the salientian morphotype, we estimated the ancestral length of the trunk region of batrachians under different hypotheses of lissamphibian relationships and divergence times. Most of our results suggest that some trunk reduction took place before the divergence of caudates and salientians (presumably in the Permian), and that the trunk of Triadobatrachusmostly reflects this ancestral condition. Thus, trunk reduction possibly preceded the anteroposterior elongation of the ilia and the shortening of the tail seen in Triadobatrachus. We also provide an updated review of the data relevant for the use of Triadobatrachusas a calibration constraint in molecular divergence age analyses that meets recently-suggested standards.

Creature Discomfort

Fauna-criticism, Ethics and the Representation of Animals in Spanish American Fiction and Poetry


Scott M. DeVries

In Creature Discomfort: Fauna-criticism, Ethics, and the Representation of Animals in Spanish American Fiction and Poetry, Scott M. DeVries uncovers a tradition in Spanish American literature where animal-ethical representations anticipate many of the most pressing concerns from present debates in animal studies. The author documents moments from the corpus that articulate long-standing positions such as a defense of animal rights or advocacy for liberationism, that engage in literary philosophical meditations concerning mind theory and animal sentience, and that anticipate current ideas from Critical Animal Studies including the rejection of hierarchical differentiations between the categories human and nonhuman.

Creature Discomfort innovates the notion of “fauna-criticism” as a new literary approach within animal studies; this kind of analysis emphasizes the reframing of literary history to expound animal ethical positions from literary texts, both those that have been considered canonical as well as those that have long been neglected. In this study, DeVries employs fauna-criticism to examine nonhuman sentience, animal interiority, and other ethical issues such as the livestock and pet industries, circuses, zoos, hunting, and species extinction in fictional narrative and poetry from the nineteenth century, modernista, Regional, indigenista, and contemporary periods of Spanish American literature.

Evolution and Human Culture

Texts and Contexts


Gregory F. Tague

Evolution and Human Culture argues that values, beliefs, and practices are expressions of individual and shared moral sentiments. Much of our cultural production stems from what in early hominins was a caring tendency, both the care to share and a self-care to challenge others. Topics cover prehistory, mind, biology, morality, comparative primatology, art, and aesthetics. The book is valuable to students and scholars in the arts, including moral philosophers, who would benefit from reading about scientific developments that impact their fields. For biologists and social scientists the book provides a window into how scientific research contributes to understanding the arts and humanities. The take-home point is that culture does not transcend nature; rather, culture is an evolved moral behavior.

Witold Morek, Piotr Gąsiorek, Daniel Stec, Brian Blagden and Łukasz Michalczyk

Edited by A. Minelli

In this paper we describe a new apochelan species, Milnesium variefidumsp. nov. from Scotland and provide novel morphological and molecular data for Milnesium berladnicorumCiobanu et al., 2014. The new species differs from the most similar M. berladnicorumby the presence of developmental dimorphism in claw configuration, absent or weakly developed cuticular bars under claws I-III, a different arrangement of cuticular pseudoplates, and by differences in the sequences of three nuclear DNA fragments: 18S rRNA (p-distance: 0.6%), 28S rRNA (2.0%), ITS-2 (9.3%), and on mitochondrial gene COI (12.4%). Although ontogenetic claw configuration change was suspected to occur in some Milnesiumspecies, we are the first to document it through the combined use of traditional, molecular and experimental methodologies. We discuss the implications of the observed phenomenon for the taxonomy of the genus and propose a new diagnostic key to all Milnesiumspecies described up to the end of 2015. We also review other traits used for species differentiation in the genus and offer recommendations to improve the quality of future descriptions as well as suggest a need for integrative redescriptions of the known species. Finally, we propose to suppress M. dujiangensisand M. tardigradum trispinosumand suggest that M. alpigenumand M. quadrifidumare valid species that require thorough redescriptions.

Frederik H. Mollen, Barry W.M. van Bakel and John W.M. Jagt

Edited by R. Vonk

A detailed redescription of a chondrocranium from the basal Boom Clay Formation (Rupelian, Upper Oligocene) at the SVK clay pit, Sint-Niklaas (province of Oost-Vlaanderen, Belgium), previously assigned to the sawshark Pristiophorus rupeliensis, is presented. The chondrocranium is re-identified as that of an angel shark (Squatinidae), based on comparative anatomy of extant Squatina, inclusive of CT scans of Squatina africana, S. australis, S. dumeril, S. guggenheimand S. squatina, with different geographic distributions and representing all four angel shark clades as defined in a previous molecular study. Differential characters for chondrocrania listed in earlier accounts to discriminate angel shark species from the southwest Atlantic proved to be even more revealing when comparing angel sharks from different regions/clades. Despite this wide interspecific variation, the fossil chondrocranium compares well with modern Squatina, but differs in having a UUU-shaped ventral margin of the occipital region and rounded margins of the upper postorbital processes. The distal expansion of the upper postorbital processes present in modern species has not yet been observed in extinct squatinoids and might constitute a derived character for modern representatives only. Angel shark teeth and vertebrae are well known from the same basal deposit at the SVK clay pit, but Cenozoic squatinid taxonomy remains problematic. It is here discussed in detail for the Oligocene taxa S. angeloides, S. rupeliensisand S. beyrichi. For the time being, all SVK material is left in open nomenclature and referred to as Squatinasp.

Fabio Scarpa, Piero Cossu, Tiziana Lai, Daria Sanna, Marco Curini-Galletti and Marco Casu

Edited by R. Sluys

Given the pending biodiversity crisis, species delimitation is a critically important task in conservation biology, but its efficacy based on single lines of evidence has been questioned as it may not accurately reflect species limits and relationships. Hence, the use of multiple lines of evidence has been portrayed as a means to overcome identification issues arising from gene/species tree discordance, morphological convergence or recent adaptive radiations. Here, the integrative taxonomic approach has been used to address the study of the Monocelis lineataspecies complex. The taxonomic resolution of the complex is challenging, as the species lacks sclerotised copulatory structures, which as a rule of thumb aid identification in Proseriata. Eighteen populations, which encompass most of the geographic range of the complex, were studied using morphology, karyology, crossbreeding experiments and molecular analysis. These different markers provided evidence of four (karyology) to eight (morphology) discrete entities, whereas crossings showed various degrees of intersterility among the tested populations. Molecular species delimitation revealed a different number of candidate species, spanning from five (ABGD and K/θ) to 11 (GMYC). Such incongruences reflect the multifaceted evolutionary history of M. lineata s.l.and hamper the full taxonomic resolution of the complex. However, two candidate species were consistently validated by all of the markers and are described as new species: Monocelis algicolanov. sp. and M. exquisitanov. sp. The latter species appear to have a restricted distribution, and the possibility that meiofaunal taxa may be of conservation concern is discussed.

Emilia Rota, Svante Martinsson, Marco Bartoli, Anneke Beylich, Ulfert Graefe, Alex Laini, Mark J. Wetzel and Christer Erséus

Edited by R. Sluys

We analysed samples of Sparganophilus taken at the corners of its distribution area in Europe (UK, Germany and Italy). No mitochondrial genetic divergence within and amongst them was found, neither in COI nor in 16S. Further, the COI haplotype was also identical to two sequences from Ontario, Canada in the Barcoding of Life Data System (BOLD) database. Our European COI and 16S sequences showed only minimal differentiation (only 1 or 2 substitutions) from specimens newly collected in Illinois and Washington states (USA), as well as from a COI haplotype from Tennessee (USA) in BOLD. An additional COI haplotype from Illinois (found in BOLD) is 2.1% different from the other haplotypes but clearly belongs to the same lineage of Sparganophilus . This geographically broad but genetically compact group fits the morphological diagnosis of S. tamesis Benham, 1892 as revised by Jamieson (1971) and is seen as evidence that all European populations 1) belong to the same species, 2) derive from a recent introduction, 3) are conspecific with the most widespread species of Sparganophilus in North America, and that 4) S. tamesis is a senior synonym of S. eiseni Smith, 1895. The single European haplotype does not refute the possibility of its spread from a single introduced source population.

V. Deepak, Varad B. Giri, Mohammad Asif, Sushil Kumar Dutta, Raju Vyas, Amod M. Zambre, Harshal Bhosale and K. Praveen Karanth

Edited by J. van Rooijen

We revise the taxonomy of the agamid genus Sitana Cuvier, 1829, a widely distributed terrestrial lizard from the Indian subcontinent based on detailed comparative analyses of external morphology, osteology and molecular data. We sampled 81 locations spread over 160,000 km2 in Peninsular India including type localities, which represented two known and five previously undescribed species. Based on general similarity in body shape and dewlap all species were hitherto identified as members of the genus Sitana. However, Sitana deccanensis and two other morphotypes, which are endemic to north Karnataka and Maharashtra in Peninsular India, are very distinct from the rest of the known members of the genus Sitana based on their external morphology and osteology. Moreover, members of this distinct morphological group were monophyletic in the molecular tree, and this clade (clade 1) was sister to two well-supported clades (2 and 3) constituting the rest of the Sitana . The interclade genetic divergence in mtDNA between clade 1 and clades 2 and 3 was 21-23%, whereas clade 2 and clade 3 exhibited 14- 16% genetic divergence. Thus, we designate a new genus name “Sarada ” gen. nov. for species represented in Clade 1, which also includes the recently resurrected Sitana deccanensis . We describe two new species in Sarada gen. nov. and three new species in Sitana . Similarity in the dewlap of Sitana and Sarada gen. nov. is attributed to similar function (sexual signaling) and similarity in body shape is attributed to a similar terrestrial life style and/or common ancestry.


Manfred Jäch, Ján Kodada, Michaela Brojer, William D. Shepard and Fedor Čiampor Jr.

Coleoptera: Elmidae and Protelmidae contains a complete list of subfamilies, tribes, subtribes, genera, subgenera, species and subspecies, and their synonyms described before 2015. Protelmidae are here elevated from tribal rank to family rank. Other new nomenclatorial and taxonomic acts include a new substitute name, seven new generic and specific synonymies, four new combinations, two designations of type species and eight mandatory corrections of incorrect original spellings. Detailed information about the geographical distribution of each species is provided. This catalogue includes extant taxa (147 genera and 1497 species of Elmidae, four genera and six species of Protelmidae) and fossil taxa (two genera and five species of Elmidae). It is the first world catalogue of Elmidae published since 1910. Unavailable names are listed as well. Detailed explanations are provided concerning the availability and correct spelling of taxa names, correct identity and spelling of author names, correct publication dates, and correct type localities.