Subspecies Recognition Research Articles

Bioacoustic Detection of Wolves: Identifying Subspecies and Individuals by Howls.

Wolves (Canis lupus) are generally monitored by visual observations, camera traps, and DNA traces. In this study, we evaluated acoustic monitoring of wolf howls as a method for monitoring wolves, which may permit detection of wolves across longer distances than that permitted by camera traps. We analyzed acoustic data of wolves' howls collected from both wild and captive ones. The analysis focused on individual and subspecies recognition. Furthermore, we aimed to determine the usefulness of acoustic monitoring in the field given the limited data for Eurasian wolves. We analyzed 170 howls from 16 individual wolves from 3 subspecies: Arctic (Canis lupus arctos), Eurasian (C. l. lupus), and Northwestern wolves (C. l. occidentalis). Variables from the fundamental frequency (f0) (lowest frequency band of a sound signal) were extracted and used in discriminant analysis, classification matrix, and pairwise post-hoc Hotelling test. The results indicated that Arctic and Eurasian wolves had subspecies identifiable calls, while Northwestern wolves did not, though this sample size was small. Identification on an individual level was successful for all subspecies. Individuals were correctly classified with 80%-100% accuracy, using discriminant function analysis. Our findings suggest acoustic monitoring could be a valuable and cost-effective tool that complements camera traps, by improving long-distance detection of wolves.

Conservation genomics and systematics of a near-extinct island radiation.

Conservation benefits from incorporating genomics to explore the impacts of population declines, inbreeding, loss of genetic variation and hybridization. Here we use the near-extinct Mariana Islands reedwarbler radiation to showcase how ancient DNA approaches can allow insights into the population dynamics of extinct species and threatened populations for which historical museum specimens or material with low DNA yield (e.g., scats, feathers) are the only sources for DNA. Despite their having paraphyletic mitochondrial DNA (mtDNA), nuclear single nucleotide polymorphisms (SNPs) support the distinctiveness of critically endangered Acrocephalus hiwae and the other three species in the radiation that went extinct between the 1960s and 1990s. Two extinct species, A. yamashinae and A. luscinius, were deeply divergent from each other and from a third less differentiated lineage containing A. hiwae and extinct A. nijoi. Both mtDNA and SNPs suggest that the two isolated populations of A. hiwae from Saipan and Alamagan Islands are sufficiently distinct to warrant subspecies recognition and separate conservation management. We detected no significant differences in genetic diversity or inbreeding between Saipan and Alamagan, nor strong signatures of geographical structuring within either island. However, the implications of possible signatures of inbreeding in both Saipan and Alamagan, and long-term population declines in A. hiwae that pre-date modern anthropogenic threats require further study with denser population sampling. Our study highlights the value that conservation genomics studies of island radiations have as windows onto the possible future for the world's biota as climate change and habitat destruction increasingly fragment their ranges and contribute to rapid declines in population abundances.

Lack of subspecies-recognition in breeding Barn Swallows (Hirundo rustica transitiva)

Assortative social interactions based on (sub)species recognition can be a driving force in speciation processes. To determine whether breeding Barn Swallows Hirundo rustica transitiva in Israel behave differentially towards members of their own subspecies, relative to a different, transient subspecies H. r. rustica and two sympatrically breeding species (Sand Martin Riparia riparia and House Sparrow Passer domesticus), we conducted a territory intrusion experiment near active nests using taxidermy models. Females responded less to the models than males, and the patterns of the recorded behavioral response traits co-varied statistically with sub- or species identity of the models, but none showed patterns of response selectivity for con(sub)specific model types only. These results do not support a role for subspecies recognition in the territorial intrusion responses of H. r. transitiva.

Phylogeography, morphology and ecological niche modelling to explore the evolutionary history of Azure-crowned Hummingbird (Amazilia cyanocephala, Trochilidae) in Mesoamerica

The Pleistocene climate cycles and mountain uplift both affected the diversification of taxa in Mesoamerica. Yet, phylogeographic breaks and demographic responses of co-distributed taxa are not congruent. Investigation of more species with widespread distributions across Mexico and Central America will help shed light on how Earth historical events shaped biodiversity in this region. The Azure-crowned Hummingbird (Amazilia cyanocephala) is widely distributed from eastern Mexico into Central America and has been divided into three subspecies by morphology and geographic isolation. To investigate the evolutionary history of the species, we sequenced a non-coding mtDNA fragment along with a nuclear marker across its distribution, and analysed morphological and ecological variation and distributional projections derived from ecological niche modelling. Overall, mtDNA supported genetic differentiation among A. cyanocephala populations, which generally supports the existence of four lineages: two A. c. cyanocephala lineages separated by the Isthmus of Tehuantepec (east and west), A. c. guatemalensis populations separated by the Motagua-Polochic-Jocotan fault system, and the isolated A. c. chlorostephana in Honduras and Nicaragua. However, morphological data were not consistent with this phylogeographic result. The potential distribution of suitable habitat for A. cyanocephala was expanded during the Last Glacial Maximum and more contracted and fragmented during the Last Inter-Glacial and the Present. These models are consistent with a preglacial demographic expansion in A. c. cyanocephala. Mitochondrial genetic diversity and current precipitation seasonality were negatively correlated when analysing all populations. We suggest that isolation and habitat differences have both played a role in the recent diversification of A. cyanocephala. Inferences about the demographic consequences of isolation and subspecies recognition await further study.

Subspecies description rates are higher in morphologically complex land snails

AbstractSubspecies richness (average number of subspecies per species = SSP/SP) has traditionally been explained by niche breadth, characteristics of the preferred habitat (“habitat hypothesis”), dispersal ability, range size (“range size hypothesis”), etc. However, in several invertebrate groups, there are striking differences between SSP/SP across families, which cannot be explained using classic arguments, and we might assume some hidden factor in the background. For example, some families of terrestrial molluscs, that possess conspicuously more subspecies than others, have higher degree of morphological complexity (shell size, number of diagnosable shell characters), for which we propose the “artefact hypothesis.” According to this, morphological complexity plays an important role in taxonomists' subspecies recognition. We used shell size, number of characters mentioned in species descriptions and area extent in six European land snail families, and habitat characteristics (rock‐dwelling vs. non‐rock‐dwelling) in one largest European snail families (Clausiliidae) to test our hypothesis. We found that all three factors (range size, morphological complexity and habitat type) correlated positively with the SSP/SP ratio. Although the generality of our findings has to be tested in other invertebrate phyla, we provide examples of striking differences between SSP/SP ratios in butterflies (Lepidoptera) and beetles (Coleoptera) as well. Our findings imply that researchers (a) should use the number of SSP more carefully in comprehensive studies, and (b) should maintain and use the rank subspecies in order to provide a stronger status for the species rank, and avoid taxonomic inflation. Furthermore, the subspecies rank might be more useful for “cryptic species” (clades that are indistinguishable morphologically) than species.

Both Environment and Genetic Makeup Influence Sexual Behavior of House Mouse

We examined changes in behavioral elements of Mus musculus musculus and Mus musculus domesticus while encountering with conspecific, allospecific animals. To do this, pure (senso lato) M. m. musculus and pure (senso lato) M. m. domesticus were obtained from the north east and the west of Iran, respectively, and mating experiments were conducted for both sub. Our study of the behavioral elements of opposite sex as the mate preference factors, within and between the two subspecies, showed that females represent more specific behavior than males in all the test groups. Also both females and males showed non-social behavior during inter-subspecies opposite sex encounters, but the aggressive behavior was just observed more when females encountered the other subspecies males. Although further studies on recognition signals (soiled bedding and urine) as a subspecies recognition system will be necessary to show inter-subspecific behavioral differences that may shape pre-mating isolation between subspecies in Iran.

Do changes in gene expression contribute to sexual isolation and reinforcement in the house mouse?

Expression divergence, rather than sequence divergence, has been shown to be important in speciation, particularly in the early stages of divergence of traits involved in reproductive isolation. In the two European subspecies of house mice, Mus musculus musculus and Mus musculus domesticus, earlier studies have demonstrated olfactory-based assortative mate preference in populations close to their hybrid zone. It has been suggested that this behaviour evolved following the recent secondary contact between the two taxa (~3,000years ago) in response to selection against hybridization. To test for a role of changes in gene expression in the observed behavioural shift, we conducted a RNA sequencing experiment on mouse vomeronasal organs. Key candidate genes for pheromone-based subspecies recognition, the vomeronasal receptors, are expressed in these organs. Overall patterns of gene expression varied significantly between samples from the two subspecies, with a large number of differentially expressed genes between the two taxa. In contrast, only ~200 genes were found repeatedly differentially expressed between populations within M.m.musculus that did or did not display assortative mate preferences (close to or more distant from the hybrid zone, respectively), with an overrepresentation of genes belonging to vomeronasal receptor family 2. These receptors are known to play a key role in recognition of chemical cues that handle information about genetic identity. Interestingly, four of five of these differentially expressed receptors belong to the same phylogenetic cluster, suggesting specialization of a group of closely related receptors in the recognition of odorant signals that may allow subspecies recognition and assortative mating.

Does competitive interaction drive species recognition in a house mouse secondary contact zone?

Miscommunication may induce a high risk of unnecessary escalated fights between competitors (populations to species), resulting in selection favoring signal divergence through agonistic character displacement (ACD). When signals allowing discrimination between competitors are also involved in mate recognition, ACD could explain reproductive character displacement (RCD). We tested interference competition between males as a potential driver of RCD (here, subspecies recognition) in a secondary contact zone between two mouse subspecies (Mus musculus musculus and Mus musculus domesticus) displaying asymmetric dominance. Since such asymmetry could create a conflict between subspecies (compatibility) and quality (dominance) recognition in the contact zone, we tested for geographic variation in female preference for dominant males in the subordinate subspecies, musculus. We assessed competition between males and tested ACD during dyadic encounters comparing behavior displayed during trials between heterosubspecifics originating from populations close to the secondary contact (“contact”) and further away (“allopatric”). We also compared behavior of contact versus allopatric males during homosubspecific versus heterosubspecific trials to test whether subspecies discrimination evolved under competitive interference. Although domesticus dominated most heterosubspecific trials regardless of geographic origin, agonistic behavior was more marked (i.e., lower attack latencies) during contact than allopatric encounters, suggesting that ACD occurred. Comparing behavior during homosubspecific and heterosubspecific encounters, only allopatric musculus displayed differences, that is, higher attack latencies toward heterosubspecifics, indicating that discrimination between competitors did not evolve with ACD. Finally, although allopatric musculus females seemed to prefer dominant males, their contact counterparts did not, suggesting that “compatibility” may have outweighed “quality” under a risk of hybridization.

Biogeography and designatable units of Bombus occidentalis Greene and B. terricola Kirby (Hymenoptera: Apidae) with implications for conservation status assessments

Conservation action for species of concern requires that “designatable units” (e.g., species, subspecies, geographic races, genetically distinct forms) are clearly defined, or that the species complex is treated as a whole. Several species of bumble bee are currently threatened, and some of these have cryptic colouration (resembling other species), or form complexes that vary considerably in colour patterning. Here we address the taxonomy and distribution of Bombus occidentalis Greene and B. terricola Kirby, both of which are currently of conservation concern in North America. Bombus occidentalis includes two apparently monophyletic groups of COI barcode haplotypes (recently considered as subspecies) with ranges mostly separated by that of their sister species, B. terricola. The southern B. o. occidentalis ranges throughout the western United States and into western Canada from southern Saskatchewan and Alberta, and throughout British Columbia north to ca. 55°N; the northern B. o. mckayi Ashmead, is restricted to north of this in British Columbia, westernmost Northwest Territories, Yukon Territory and Alaska. Bombus o. mckayi exists, as far as is known, only with a “banded” colour pattern. By contrast, B. o.occidentalis occurs in both banded and non-banded colour patterns, although the southern banded colour pattern is geographically isolated from the northern subspecies. Bombus o. occidentalis has declined throughout its range, perhaps due in part to exposure to novel parasites. Despite having similar levels of parasitism (ca. 40 %) as the southern subspecies, B. o. mckayi appears to have stable populations at present. There is therefore compelling evidence that the two subspecies should be distinguished for conservation and management purposes. We present the evidence for their distinction and provide tools for subspecies recognition.

Ultrasonic Vocalizations of Male Mice Differ among Species and Females Show Assortative Preferences for Male Calls.

Male house mice (Mus musculus) emit ultrasonic vocalizations (USVs) during courtship, which attract females, and we aimed to test whether females use these vocalizations for species or subspecies recognition of potential mates. We recorded courtship USVs of males from different Mus species, Mus musculus subspecies, and populations (F1 offspring of wild-caught Mus musculus musculus, Mus musculus domesticus (and F1 hybrid crosses), and Mus spicilegus), and we conducted playback experiments to measure female preferences for male USVs. Male vocalizations contained at least seven distinct syllable types, whose frequency of occurrence varied among species, subspecies, and populations. Detailed analyses of multiple common syllable types indicated that Mus musculus and Mus spicilegus could be discriminated based on spectral and temporal characteristics of their vocalizations, and populations of Mus musculus were also distinctive regardless of the classification model used. Females were able to discriminate USVs from different species, and showed assortative preferences for conspecific males. We found no evidence that females discriminate USVs of males from a different subspecies or separate populations of the same species, even though our spectral analyses identified acoustic features that differ between species, subspecies, and populations of the same species. Our results provide the first comparison of USVs between Mus species or between Mus musculus subspecies, and the first evidence that male USVs potentially facilitate species recognition.

Diagnosability and description of a new subspecies of Indo-Pacific humpback dolphin, Sousa chinensis (Osbeck, 1765), from the Taiwan Strait.

Subspecies recognition can affect how people (scientists and non-scientists alike) view organisms and thus has important implications for research on, as well as the conservation of, these entities. Recently, a small group of Indo-Pacific humpback dolphins was discovered inhabiting the waters off central western Taiwan. This geographically isolated population possesses pigmentation patterns that are subtly, but noticeably, different from their nearest conspecifics in the neighbouring waters of the Jiulong River Estuary and Pearl River Estuary of mainland China. Due to this population's low and declining numbers and the numerous threats it faces, it was assessed as critically endangered by the Red List of Threatened Species of the International Union for Conservation of Nature. The purpose of this study is to examine the degree of differentiation of the Taiwanese population to determine if subspecies recognition iswarranted. Analysis of the degree of differentiation in pigmentation patterns revealed nearly non-overlapping distributions between dolphins from Taiwanese waters and those from the Jiulong River + Pearl River estuaries of mainland China (the nearest known populations). The Taiwanese dolphins were clearly diagnosable from those of the Jiulong River + Pearl Riverestuaries under the most commonly accepted '75%rule' for subspecies delimitation (with 94% of one group being separable from 99+% of the other). Evidence of geographical isolation and behavioural differences also provided additional support for the distinctiveness of the Taiwanese dolphins. Together, the evidence strongly demonstrated that the Taiwanese humpback dolphin population is differentiated at the subspecies level and on an evolutionary trajectory that is independent from that of dolphins from adjacent waters of mainland China (i.e. Jiulong River + Pearl River estuaries). As a result, the taxonomy of Sousa chinensis was revised to include two subspecies: the Taiwanese humpback dolphin, Sousa chinensis taiwanensis subsp. nov., and the Chinese humpback dolphin, Sousa chinensis chinensis (the nominotypical subspecies). These subspecies are described, and the holotype and paratype specimens for S. c. taiwanensis are established.

A review of subspecies recognition in polydesmidan millipedes (Diplopoda) with a revision of the subspecies of Euryuridae (Xystodesmoidea)

Jorgensen, M. C., Sierwald, P. & Mason‐Gamer, R. J. (2012). A review of subspecies recognition in polydesmidan millipedes (Diplopoda) with a revision of the subspecies of Euryuridae (Xystodesmoidea). —Zoologica Scripta, 42, 317–326.Taxonomic subspecies recognition is controversial due to the lack of an objective definition of the concept and inconsistent use of the category. The practice of designating subspecies is assessed here through a thorough review of 244 subspecies designations of polydesmidan millipedes over a 50‐year period. The survey focuses on the justification given for subspecies recognition, the amount of data available for the designation, the handling of nominate subspecies and the criteria used for diagnosis. We address several problematic issues and provide suggestions to enhance future work. Three examples of subdivided species from the Euryuridae (Polydesmida) are presented in detail with some taxonomic revision. Euryurus leachii leachii and E. leachii fraternus are synonymized, and the subspecific epithets are discarded. Auturus erythropygos erythropygos and A. erythropygos becki are returned to full species rank.

Differential response of males of a subarctic population of Bluethroat Luscinina svecica svecica to playbacks of their own and foreign subspecies

Differences in vocalization between subspecies of songbirds may influence intraspecific interactions including mate choice and territorial behavior, and contribute to their further divergence by strengthening pre-mating reproductive isolation. We have recently shown that songs of two Bluethroat subspecies, Luscinia svecica svecia and L. s. cyanecula, differ in spectral characteristics and structure. Here, in a first experimental test of whether these differences are biologically relevant, we compared responses of L. s. svecica males to both types of songs. Altogether 19 males from a subartic population in Abisko, Sweden, were exposed to playbacks of songs from the two subspecies, accompanied by display of a stuffed Bluethroat lacking subspecies-specific colour marks. These stimuli were presented in random order, separated by over 1 h. We evaluated two largely uncorrelated components of male responses: non-vocal territorial behavior, and singing activity. A significant difference in reaction to subspecies playbacks was observed in aggressive behavior: independently of the song order, males generally responded more strongly to playback of their own subspecies. In contrast, playback order rather than subspecies identity significantly influenced the singing activity, which usually increased in the second trial. A more detailed analysis nevertheless suggested that vocal responses of birds exposed first to playback of their own subspecies were stimulated in the subsequent trial but not vice versa. Our results show that Bluethroats clearly discriminate the two song types, indicating the potential for subspecies recognition. Further experimental work is needed to assess the general validity of these patterns.

Song divergence of two Bluethroat subspecies (Luscinia s. svecica and L. s. cyanecula)

As birds adapt to changing biotic and abiotic environmental conditions, parameters of their acoustic signals are modified by selective pressure for efficient communication. Such changes in song features may result in acoustic divergence among populations of the same species. We studied the divergence of songs in four populations of two genetically distinct subspecies of the Bluethroat Luscinia svecica – L. s. svecica and L. s. cyanecula. Males of svecica, occupying tundra habitats, produced less diverse songs with lower fre-quencies than cyanecula males occurring in several types of habitat. The differences in frequency parameters may be related to larger body size of the former subspecies; how-ever, habitat associations predict an opposite relationship. Contrary to expectations, rep-ertoire size did not significantly differ between the two subspecies, although cyanecula inhabits habitats with higher density and diversity of songbirds (which it may imitate). Temporal song parameters (song length, pause length, and song rate) differed signifi-cantly among populations but not between subspecies. Differences in song characteristics might be important in subspecies recognition, and may therefore contribute to the mainte-nance of subspecific differences or even influence the speciation process.

A candidate subspecies discrimination system involving a vomeronasal receptor gene with different alleles fixed in M. m. domesticus and M. m. musculus

Assortative mating, a potentially efficient prezygotic reproductive barrier, may prevent loss of genetic potential by avoiding the production of unfit hybrids (i.e., because of hybrid infertility or hybrid breakdown) that occur at regions of secondary contact between incipient species. In the case of the mouse hybrid zone, where two subspecies of Mus musculus (M. m. domesticus and M. m. musculus) meet and exchange genes to a limited extent, assortative mating requires a means of subspecies recognition. We based the work reported here on the hypothesis that, if there is a pheromone sufficiently diverged between M. m. domesticus and M. m. musculus to mediate subspecies recognition, then that process must also require a specific receptor(s), also sufficiently diverged between the subspecies, to receive the signal and elicit an assortative mating response. We studied the mouse V1R genes, which encode a large family of receptors in the vomeronasal organ (VNO), by screening Perlegen SNP data and identified one, Vmn1r67, with 24 fixed SNP differences most of which (15/24) are nonsynonymous nucleotide substitutions between M. m. domesticus and M. m. musculus. We observed substantial linkage disequilibrium (LD) between Vmn1r67 and Abpa27, a mouse salivary androgen-binding protein gene that encodes a proteinaceous pheromone (ABP) capable of mediating assortative mating, perhaps in conjunction with its bound small lipophilic ligand. The LD we observed is likely a case of association rather than residual physical linkage from a very recent selective sweep, because an intervening gene, Vmn1r71, shows significant intra(sub)specific polymorphism but no inter(sub)specific divergence in its nucleotide sequence. We discuss alternative explanations of these observations, for example that Abpa27 and Vmn1r67 are coevolving as signal and receptor to reinforce subspecies hybridization barriers or that the unusually divergent Vmn1r67 allele was not a product of fast positive selection, but was derived from an introgressed allele, possibly from Mus spretus.

A Candidate Subspecies Discrimination System Involving a Vomeronasal Receptor Gene with Different Alleles Fixed in M. m. domesticus and M. m. musculus

Assortative mating, a potentially efficient prezygotic reproductive barrier, may prevent loss of genetic potential by avoiding the production of unfit hybrids (i.e., because of hybrid infertility or hybrid breakdown) that occur at regions of secondary contact between incipient species. In the case of the mouse hybrid zone, where two subspecies of Mus musculus (M. m. domesticus and M. m. musculus) meet and exchange genes to a limited extent, assortative mating requires a means of subspecies recognition. We based the work reported here on the hypothesis that, if there is a pheromone sufficiently diverged between M. m. domesticus and M. m. musculus to mediate subspecies recognition, then that process must also require a specific receptor(s), also sufficiently diverged between the subspecies, to receive the signal and elicit an assortative mating response. We studied the mouse V1R genes, which encode a large family of receptors in the vomeronasal organ (VNO), by screening Perlegen SNP data and identified one, Vmn1r67, with 24 fixed SNP differences most of which (15/24) are nonsynonymous nucleotide substitutions between M. m. domesticus and M. m. musculus. We observed substantial linkage disequilibrium (LD) between Vmn1r67 and Abpa27, a mouse salivary androgen-binding protein gene that encodes a proteinaceous pheromone (ABP) capable of mediating assortative mating, perhaps in conjunction with its bound small lipophilic ligand. The LD we observed is likely a case of association rather than residual physical linkage from a very recent selective sweep, because an intervening gene, Vmn1r71, shows significant intra(sub)specific polymorphism but no inter(sub)specific divergence in its nucleotide sequence. We discuss alternative explanations of these observations, for example that Abpa27 and Vmn1r67 are coevolving as signal and receptor to reinforce subspecies hybridization barriers or that the unusually divergent Vmn1r67 allele was not a product of fast positive selection, but was derived from an introgressed allele, possibly from Mus spretus.

A host species‐informative internal control for molecular assessment of African swine fever virus infection rates in the African sylvatic cycle Ornithodoros vector

African swine fever virus (ASFV) infection in adult Ornithodoros porcinus (Murry 1877, sensuWalton 1979) ticks collected from warthog burrows in southern and East Africa was assessed using a duplex genomic amplification approach that is informative with respect to the invertebrate host species and infecting sylvatic cycle virus. DNA extracted from individual ticks was used as template for the simultaneous amplification of a C-terminal 478-bp ASFV p72 gene region and a approximately 313-bp fragment of the tick mitochondrial 16S rRNA gene, under optimized reaction conditions. Within-warthog burrow infection rates ranged from 0% to 43% using this approach, and phylogenetic analysis of 16S gene sequences revealed the presence of three geographically discrete O. porcinus lineages, but no support for subspecies recognition. False negatives are precluded by the inclusion of host species-informative primers that ensure the DNA integrity of cytoplasmically located genome extracts. In addition, infection rate estimates are further improved as false positives arising from carry-over contamination when performing a two-step nested polymerase chain reaction are negated by the one-step approach. Phylogenetic comparison of full-length virus gene sequences with the partial C-terminal p72 gene target confirmed the epidemiological utility of the latter in a sylvatic setting. The method is therefore of particular value in studies assessing the prevalence and diversity of ASFV in relation to the African sylvatic tick vector and holds potential for investigating the role of alternative tick species in virus maintenance and transmission.

Problems of subspecies recognition in freshwater cyclopoid copepods

Summary The systematics of freshwater copepods has been considered from a historical viewpoint. These arthropods are characterized by considerable phenotypical plasticity, many sibling-species, polymorphism and polyphenism. The taxonomy within freshwater copepods is mostly based upon the morphology of the animals and the splitting of a number of species into subspecies is often based on very scarce material. Seldom have taxonomists working on freshwater copepods used adequate taxonomic theory or taken advantage of the tools of modem taxonomy. A systematic analysis of the species Cyclops abyssorum has been performed. Selected populations within the species range have been measured and compared with the topotypical population. Geographically neighbouring populations may differ substantially, while more remote populations may bear a greater resemblance to each other. Analysis of the life cycle strategies did not correlate with the morphological analysis. C. abyssorum does not satisfy modern taxonomical rules for splitting into subspecies. Local or geographically restricted populations will be denoted by using the binomen and the locality. Zusammenfassung Probleme der Unterscheidung von Unterarten bei cyclopoiden Süβwassercopepoden Die Systematik der Süßwassercopepoden wird historisch betrachtet. Diese Arthropoden zeichnen sich durch eine bemerkenswerte Plastizität, viele Zwillingsarten, Polymorphismen und Polyphaenismen aus. Der Taxonomie der Süßwassercopepoden wird im allgemeinen die Morpholoie zugrunde gelegt, Aufsplitterungen in Unterarten beruhen meist auf einem sehr kleinen Material. Selten benutzen Taxo-nomen dieser Gruppe in angemessener Weise taxonomische Theorien oder die Hilfsmittel moderner Taxonomie. Eine taxonomische Analyse der Art Cyclops abyssorurn wird vorgelegt. Ausgewählte Populationen aus dem gesamten Verbreitungsgebiet wurden vermessen und mit topotyischen Poulationen ver-glichen. Es zeigt sich, daß geographisch benachbarte Populationen sich deuich unterscheiden können, während voneinander weit endernte Populationen sich ähneln. Biologische Besonderheiten stehen mit Gestalteigenarten nicht in Korrelation. Bei Zugrundelegung moderner taxonomischer Grundsätze läßt sich C. abyssorum nicht in Unterarten aufgliedem. Lokale oder geographisch begrenzte Populationen sollten mit dem Binomen und der Lokalität gekennzeichnet werden.

Variation in C-stained chromosome regions in European hedgehogs (Insectivora, Mammalia).

European hedgehogs were studied by C- and G-banding techniques. Animals were sampled from several localities in Europe, but mainly from the Scandinavian countries. The interest was focused on the presence or absence of large autosomal C-blocks in individual chromosome pairs, and on the incidence of C-band variants. With respect to number and distribution of such blooks five different karyotypes could be distinguished in the material, three belonging to the Western hedgehogs and two to the Eastern ones. These dissimilarities were the most significative differences between karyotypes. The geographical distribution of the five karyotypes is discussed with reference to existing subspecies recognition. The large distal C-blocks appearing in 2–4 chromosome pairs were highly polymorphic in size, reflected in a high frequency of heteromorphic pairs. An attempt was made to shed some light on the problem of a continuous or discontinuous type of variation of the C-blocks and, therefore, a statistical method was applied to quantitative data. Only one sample of 22 animals from southern Sweden was regarded as informative, indicating that the size variation of the C-blocks in one chromosome pair was bi- or trimodal, and thus discontinuous. Contrarily, the size distribution of the C-blocks in the other two pairs seemed to be continuous, although their mode of variation differed slightly.

Description of a new subspecies of the crayfish Parastacus brasiliensis (Von Martens, 1869) from Sao Francisco de Paula, RS, Brazil (Decapoda, Parastacidae)

Parastacus brasiliensis promatensis subsp. n. is described from São Francisco de Paula, Rio Grande do Sul, Brazil. The animals were collected at an elevation of 850 m, representing the first record of Parastacidae at high altitude in Brazil. Subspecies recognition was performed through the analysis of morphometrical characters. Discriminant analysis was applied to compare P. brasiliensis promatensis subsp. n. with the known populations of P. brasiliensis (Von Martens, 1869) using residuals from 23 morphometric regressions (Canonical Correlation = 0.948). All variance was explained by function 1 (Pbrasiliensis promatensis centroid equal to 3.04; P. brasiliensis centroid equal to -2.78).