Conflicting molecular phylogenies of European long-eared bats (Plecotus) can be explained by cryptic diversity

Conflicting phylogenetic signals of two data sets that analyse different portions of the same molecule are unexpected and require an explanation. In the present paper we test whether (i) differential evolution of two mitochondrial genes or (ii) cryptic diversity can better explain conflicting results of two recently published molecular phylogenies on the same set of species of long-eared bats (genus Plecotus). We sequenced 1714bp of three mitochondrial regions (16S, ND1, and D-loop) of 35 Plecotus populations from 10 European countries. A likelihood ratio test revealed congruent phylogenetic signals of the three data partitions. Our phylogenetic analyses demonstrated that the existence of a previously undetected Plecotus lineage caused the incongruities of previous studies. This lineage is differentiated on the species level and lives in sympatry with its sister lineage, Plecotus auritus, in Switzerland and Northern Italy. A molecular clock indicates that all European Plecotus species are of mid or late Pliocene origin. Plecotus indet. was previously described as an intergrade between P. auritus and Plecotus austriacus since it shares morphological characters with both. It is currently known from elevations above 800 m a.s.l. in the Alps, the Dinarian Alps and the Pindos mountains in Greece. Since we could demonstrate that incongruities of two molecular analyses simply arose from the mis-identification of one lineage, we conclude that molecular phylogenetic analyses do not free systematists from a thorough inclusion of morphological and ecological data.     

Cryptic diversity in European bats

Different species of bat can be morphologically very similar. In order to estimate the amount of cryptic diversity among European bats we screened the intra- and interspecific genetic variation in 26 European vespertilionid bat species. We sequenced the DNA of subunit 1 of the mitochondrial protein NADH dehydrogenase (ND1) from several individuals of a species, which were sampled in a variety of geographical regions. A phylogeny based on the mitochondrial (mt) DNA data is in good agreement with the current classification in the family. Highly divergent mitochondrial lineages were found in two taxa, which differed in at least 11% of their ND1 sequence. The two mtDNA lineages in Plecotus austriacus correlated with the two subspecies Plecotus austriacus austriacus and Plecotus austriacus kolombatovici. The two mtDNA lineages in Myotis mystacinus were partitioned among two morphotypes. The evidence for two new bat species within Europe is discussed. Convergent adaptive evolution might have contributed to the morphological similarity among distantly related species if they occupy similar ecological niches. Closely related species may differ in their ecology but not necessarily in their morphology. On the other hand, two morphologically clearly different species (Eptesicus serotinus and Eptesicus nilssonii) were found to be genetically very similar. Neither morphological nor mitochondrial DNA sequence analysis alone can be guaranteed to identify species.     

Assessment of available anatomical characters for linking living mammals to fossil taxa in phylogenetic analyses

Analyses of living and fossil taxa are crucial for understanding biodiversity through time. The total evidence method allows living and fossil taxa to be combined in phylogenies, using molecular data for living taxa and morphological data for living and fossil taxa. With this method, substantial overlap of coded anatomical characters among living and fossil taxa is vital for accurately inferring topology. However, although molecular data for living species are widely available, scientists generating morphological data mainly focus on fossils. Therefore, there are fewer coded anatomical characters in living taxa, even in well-studied groups such as mammals. We investigated the number of coded anatomical characters available in phylogenetic matrices for living mammals and how these were phylogenetically distributed across orders. Eleven of 28 mammalian orders have less than 25% species with available characters; this has implications for the accurate placement of fossils, although the issue is less pronounced at higher taxonomic levels. In most orders, species with available characters are randomly distributed across the phylogeny, which may reduce the impact of the problem. We suggest that increased morphological data collection efforts for living taxa are needed to produce accurate total evidence phylogenies.     

Morphological definition of Eurotestudo (Testudinidae,Chelonii): First part

This work details the morphological analysis upon which the creation of the testudinid genus Eurotestudo was based. The study is presented in two parts. The present first part includes an introduction to the subject, and the morphological data with illustrations, which formed the basis for the analysis. The results and lists of cited taxa appear in the second part (Ann. Pal., 92, 4). The analysis rests on a limited number of selected characters, principally those derived from shell osteology, which are preserved in fossils as well as in living specimens. Within the palaearctic genus Testudo s.l., Eurotestudo was separated from two other inclusive genera, Agrionemys and Testudo s.s. In a cladistic analysis, Eurotestudo is distinguished by the autapomorphy of the narrowing vertebrals. Several more external characters (not derivable from fossils and thus omitted from the present cladistic analysis) also support the existence of the genus. Eurotestudo shares homoplastic characters with the two other lineages, the frequency and the evolutionary degree of which vary according to species in the two other genera. Further osteological characters and characters known only from living specimens are added to the osteological study. The characters are examined on a series of taxa chosen among the better known fossil forms from the Oligocene to the Present, Quaternary forms of the Eu. hermanni group included, and representative extant forms. Possible members of the three stems are examined. The autapomorphies of each stem are evidenced, the polarization of characters is tested and the homoplasies are detected. The hypothesis of a common origin for Testudo and Eurotestudo has best support in the present analysis.     

The classification of ferns

Intensive morphological studies have been devoted to the more primitive ferns, which represent a small minority of living species, but too little is yet known about the vast majority of other ferns, with the result that recent attempts at a natural classification show considerable differences of treatment.
The problem is complicated by convergent evolution in the characters of almost all parts of a fern plant. Not only similar soral form, but also similar frond form, types of venation, scales, etc. have been developed on different evolutionary lines.
To illustrate the nature of the problem an attempt has been made to state the probable characters of a primitive leptosporangiate fern, and the kinds of ways in which existing ferns have developed from this condition. Evolutionary change in different parts of the plant has proceeded in different ways and to different degrees in the many genera of existing ferns. Primitive characters of one kind or another are shown by a great number of ferns, along with highly advanced characters of other kinds.
Recent schemes of classification are briefly compared, and a summary is given of the author's own scheme, with notes on evolutionary trends in the various groups as he sees them.
Much more information is needed on which to establish a really satisfactory scheme. The present one is put forward in the hope that others will take up the work. With modern facilities for travel, it is to be hoped that more botanists will come to the tropics and see ferns and other too-little-known plants in their native habitats. Morphological study needs to be undertaken with an understanding of the living plant and of its environment.     

A preliminary revision of the genus Plecotus (Chiroptera, Vespertilionidae) based on genetic and morphological results

The phylogenetic relationships within the genus Plecotus were assessed using molecular as well as morphological methods. With only three species missing, our study is based on an almost comprehensive taxonomic sampling. The genetic analysis comprised 151 individuals from throughout the range. Sequences of two mitochondrial sections, parts of the 16S rRNA gene (16S) and of the control region (CR) were analysed. The morphological analysis of cranial and external characters comprised 697 individuals, including 10 holotypes and one lectotype. Data from 15 craniometric characters of 442 specimens were used in the multivariate analyses. The molecular data identified nine primary clades representing 11 species, 10 of which could be assigned to described taxa, whereas one was described as a new species, Plecotus strelkovi Spitzenberger sp. nov. The tree based on 16S revealed two major lineages, one consisting of only one primary clade restricted to the Mediterranean, the other consisting of eight primary clades representing Eurasian taxa. The morphological analysis revealed five additional species, two of them not described. Together with the recently described P. taivanus, P. sardus and P. balensis, which were not included in our analysis, the genus Plecotus comprises at least 19 more or less cryptic species. Phylogenetic and phenetic analyses resulted in similar but not completely concordant arrangements of the species. The proposed classification relies mainly on the tree based on 16S sequences. The current distribution indicates that 16 species can be linked to arboreal refugia, three to eremial refugia. We assume that speciation within the gleaning, rather slow flying long‐eared bats is due to a multitude of disruption and isolation processes within a formerly continuous range of the broad‐leaved Arcto‐Tertiary forest in which Plecotus probably originated. An exact calibrated molecular dating of the splits is not possible. The Early Oligocene age of the presumed ancestor of the Plecotini and a correlation of the molecular diversifications with palaeogeographic reconstructions suggest that the divergence of the two major lineages may have occurred already during the Middle Miocene, 14.5 Mya.     

Comparing levels of homoplasy in the primate skeleton

Hard-tissue morphological characters (bones and teeth) are a primary source of information about the evolutionary history of primates. These tissues are commonly found as isolated elements in the fossil record and studied as three separate partitions: the dentition, the cranium, and the postcranium. The relative phylogenetic utility of characters from each partition is often called into question with respect to varying amounts of homoplasy. In this paper, the consistency index (CI) was used to measure levels of homoplasy in each data partition for a sample of fossil and living primates. Sources of bias in the collection and treatment of data and in the internal structure of the data set are addressed. These biases include number of taxa, number of characters, ordering of characters, amounts of polymorphically scored or missing data, and character-state distribution. The results of this study suggest that the levels of homoplasy are very similar, though the postcranial data may be slightly less homoplastic than either the dental or cranial data.     

Mitochondrial phylogeography of the long-eared bats (Plecotus) in the Mediterranean Palaearctic and Atlantic Islands

Long-eared bats of the genus Plecotus are widespread and common over most of the western Palaearctic. Based on recent molecular evidence, they proved to represent a complex of several cryptic species, with three new species being described from Europe in 2002. Evolutionary relationships among the different lineages are still fragmentary because of the limited geographic coverage of previous studies. Here we analyze Plecotus mitochondrial DNA sequences from the entire Mediterranean region and Atlantic Islands. Phylogenetic reconstructions group these western Palaearctic Plecotus into two major clades which split at least 5 Myr ago and that are each subdivided into further subgroups. An 'auritus group' includes the traditional P. auritus species and its sister taxon P. macrobullaris (=P. alpinus) plus related specimens from the Middle East. P. auritus and P. macrobullaris have broadly overlapping distributions in Europe, although the latter is apparently more restricted to mountain ranges. The other major clade, the 'austriacus group,' includes the European species P. austriacus and at least two other related taxa from North Africa (including P. teneriffae from the Canary Islands), the Balkans and Anatolia (P. kolombatovici). The sister species of this 'austriacus group' is P. balensis, an Ethiopian endemic. Phylogenetic reconstructions further suggest that P. austriacus reached Madeira during its relatively recent westward expansion through Europe, while the Canary Islands were colonized by a North African ancestor. Although colonization of the two groups of Atlantic Islands by Plecotus bats followed very distinct routes, neither involved lineages from the 'auritus group.' Furthermore, the Strait of Gibraltar perfectly segregates the distinct lineages, which confirms its key role as a geographic barrier. This study also stresses the biogeographical importance of the Mediterranean region, and particularly of North Africa, in understanding the evolution of the western Palaearctic biotas.     

Climate Analysis of Endemic Species-A Novel Method for Quantitative Analysis of Global Climate Change Since Tertiary

There are many extant endemic plants in China, which were widely distributed in the North Hemisphere during Tertiary. The global cooling during the Tertiary caused a series of narrow distribution regions of the plants. Quaternary glaciation invaded most regions of North America and Eurasia where severe destruction was imposed onto vegetation. However, such destruction was lessened in China largely because of specific topographic and geographical and obviously, a number of other conditions accounted for an unusual refugee camp for the relics of plants in China, among which lots of endemic taxa exist. Recently, Chinese endemic species, such as Metaseqouia, Eucommia , have been employed to conduct multi-disciplinary comprehensive studies so as to analyze Tertiary climate changes quantitatively. Meanwhile, a rigorous method, i.e. climate analysis of endemic species (CAES) has come to maturation. This method is characteristic of some generality because it is supposed to be applicable to the endemic species in other regions of the world. CAES is involved in the following aspects: 1. Conduct multidisciplinary studies on living and fossil species of endemic plants and trace their evolutionary courses. 2. Compare fossil species with living one and clarify which is the nearest living relative (NLR) to fossil counterpart. 3. Fossils and their living counterparts (NLR) are supposed to have similar ecological requirements to meet their life cycles. 4. Investigate the geographic distribution of living and fossil plants within the same taxa and ascertain the dynamic changes of their distributions in geological age. 5. Analyze climate factors in the distribution of specific endemic taxa and obtain the data of climatic characters which are suitable for reconstruction of paleoclimate where fossil counterparts lived. 6. Further study the physio-ecology of living species and determinate paleoclimate where fossil counterparts lived. 7. Integrate analysis of the data from steps 4, 5 and 6, and quantitatively reconstruct the climate where fossil and living plants survive.     

The use of cladistic principles in the phylogeny and biogeography of the Lacerta

The methodology used in Phylogenetics remains one of the more interesting problems in comparative biology. Among the methods used, the cladistic is the most accurate. In the present case, among the Saurians, the characters chosen appeared early and cannot be referred to any adaptation bound to an evolutive progress. We have successively considered their distribution in the classification, their polarity and one phylogenetic hypothesis based on the Hennig's concepts. Such a work leads necessarily to systematical conclusions. For instance, we note the heterogeneity of some families. Among the Iguania, the Iguanidea are polyphyletics, the north-american forms breaking up from the other very early. Among the Scincomorpha, the Lacertilia result from two parts of different origins. This research verifies some systematic cuttings in the modern classification: among the Anguimorpha, the separation between Helodermatidea and the group, Varanidea-Lanthanotidea. The characters studied show a non-random distribution in the classification and show also some consistency with the geographical distribution of the extant Saurians. Therefore, it is important to search for the most parsimonious geographical hypothesis, which is in concern with the present data on the continental drift and generally on the history of the Earth, and which strengthens the cladograms posed from the selected characters.     

Phylogeny and biogeography of the extant species of triplespine fishes (Triacanthidae, Tetraodontiformes)

A new phylogenetic hypothesis for the living species of triplespine fishes of the Indo-Western Pacific family Triacanthidae (Tetraodontiformes, Teleostei) is proposed. A data set of 55 morphological characters (34 osteological and 21 morphometric) was constructed. A cladistic analysis of the osteological data set yielded a single most-parsimonious tree. This cladogram does not support the monophyly of one of the four genera, Tripodichthys , but Bremer values for this analysis are low. The osteological data set was then combined with a data set of 21 morphometric characters that had previously been used to diagnose the four genera. The analysis of the combined data set produced the same phylogenetic hypothesis, but with greater nodal support. The biogeographical distribution of the living species is then interpreted with the use of this new phylogenetic information.     

Detection of major genes for quantitative morphologic characteristics on family material, sib pairs and parent-child pairs, using as an example head length in man]

The detection of major genes affecting quantitative morphological characters from data of families, sib-pairs and parent-child pairs (with the example of head length in man). The major genes of human head length are defined utilizing three different combinatorical methods of maximum likelihood estimation. To prove the discriminatory power of methods, it is necessary in further research to work with empirical or simulated sets of multivariate data of which the genetical structure is known.     

定量分析第三纪以来环境变化的新方法：—特有种气候分析法

1　全球气候变化研究新动向自 2 0世纪 80年代以来 ,全球气候变化研究至今仍方兴未艾。全球气温波动、大气中温室气体丰度变化、气候格局改变、气候带迁移和降水量增减 ,都影响到社会经济的发展和人类生活质量的提高。各国政府对全球气候变化的研究 ,如气候演变的过程、短期和长期气候变化的预测和自然灾害的防治都给予极大关注和财力支持[1] 。目前全球气候变化的研究正向两个方向发展。1.1　加大全球气候变化研究时间尺度 ,从研究当今气候发展到研究地质历史时期气候变化全球气候变化 ,如大气中ＣＯ2 浓度的变化、温度的升降和降水量的变…     

定量分析第三纪以来环境变化的新方法——特有种气候分析法

There are many extant endemic plants in China, which were widelydistributed in the North Hemisphere during Tertiary. The global cooling during the Tertiary caused a series of narrow distribution regions of the plants. Quaternary glaciation invaded most regions of North America and Eurasia where severe destruction was imposed onto vegetation. However, such destruction was lessened in China largely because of specific topographic and geographical and obviously, a number of other conditions accounted for an unusual refugee camp for the relics of plants in China, among which lots of endemic taxa exist. Recently, Chinese endemic species, such as Metaseqouia, Eucommia, have been employed to conduct multi-disciplinary comprehensive studies so as to analyze Tertiary climate changes quantitatively. Meanwhile, a rigorous method, i.e. climate analysis of endemic species (CAES) has come to maturation. This method is characteristic of some generality because it is supposed to be applicable to the endemic species in other regions of the world. CAES is involved in the following aspects: 1. Conduct multidisciplinary studies on living and fossil species of endemic plants and trace their evolutionary courses. 2. Compare fossil species with living one and clarify which is the nearest living relative (NLR) to fossil counterpart. 3. Fossils and their living counterparts (NLR) are supposed to have similar ecological requirements to meet their life cycles. 4. Investigate the geographic distribution of living and fossil plants within the same taxa and ascertain the dynamic changes of their distributions in geological age. 5. Analyze climate factors in the distribution of specific endemic taxa and obtain the data of climatic characters which are suitable for reconstruction of paleoclimate where fossil counterparts lived. 6. Further study the physio-ecology of living species and determinate paleoclimate where fossil counterparts lived. 7. Integrate analysis of the data from steps 4, 5 and 6, and quantitatively reconstruct the climate where fossil and living plants survive.     

Intraspecific evolution of Canary Island Plecotine bats,based on mtDNA sequences

Island differentiation and relationships with congenerics were investigated in the endemic Canary Island bat Plecotus teneriffae, based on approximately 1 kb of mtDNA from the 16S rRNA and cytochrome b genes. P. teneriffae had closer affinities with P. austriacus than with P. auritus. Levels of differentiation between Canary Islands were quite high relative to Pipistrelle-like bats, consistent with philopatric behaviour in the Plecotus genus. Cladogenesis within P. teneriffae appears to have occurred after the emergence of the islands of El Hierro and La Palma during the Pleistocene. An intraspecific network shows that haplotypes from the younger islands of La Palma and El Hierro are connected to the Tenerife haplotype by a similarly large number of mutational steps. This suggests that they were both colonised at a similar time from the much older island of Tenerife. The other Plecotine bat species, Barbastellus barbastellus shows close affinities with B. barbastellus from mainland Spain, with levels of mtDNA divergence being comparable with intraspecific variation within other mammal species.     

Impact of increased character sampling on the phylogeny of Cetartiodactyla (Mammalia): combined analysis including fossils

The phylogenetic position of Cetacea (whales, dolphins and porpoises) is an important exemplar problem for combined data parsimony analyses because the clade is ancient and includes many well‐known and relatively complete fossil species. We combined data for 71 terminal taxa (43 extinct/28 extant) to test where Cetacea fits within Cetartiodactyla, and where various fossil hoofed mammals (e.g., ?entelodonts, “?anthracotheriids” and ?mesonychians) are positioned. We scored 635 phenotypic characters (osteology, dentition, soft tissue, behavior), approximately three times the number of characters in the last major analysis of this clade, and combined these with > 40 000 molecular characters, including new data from 10 genes. The analysis supported a topology consistent with the majority of recently published molecular studies. Cetacea was the extant sister taxon of Hippopotamidae, followed successively by Ruminantia, Suina and Camelidae. Several extinct taxa were phylogenetically unstable, upsetting resolution of the strict consensus and limiting branch support, but the positions of several key fossils were consistently resolved. The wholly extinct ?Mesonychia was more closely related to Cetacea than was any “artiodactylan.”“?Anthracotheriids” were paraphyletic, and, with the exception of one species, were more closely related to Hippopotamidae than to any other living taxon. The total evidence analysis overturned a highly nested position for Moschus supported by molecular data alone. The character partition that could be scored for the fossil taxa (osteological and dental characters) included more informative characters than most molecular partitions in our analysis, and had the fewest missing data. The osteological–dental data alone, however, did not support inclusion of cetaceans within crown “Artiodactyla.” Recently discovered ankle bones from fossil whales reinforced the monophyly of Cetartiodactyla but provided no particular evidence of derived similarities between hippopotamids and fossil cetaceans that were not shared with other “artiodactylans”. © The Willi Hennig Society 2007.     

Molecular phylogeny of North American long-eared bats (Vespertilionidae: Corynorhinus); inter- and intraspecific relationships inferred from mitochondrial and nuclear DNA sequences

The taxonomy of the North American big-eared bats, genus Corynorhinus, was revised by Handley [Handley, C.O., 1959. A revision of the American bats of the genera Euderma and Plecotus. Proc. U.S. Nat. Mus. 110, 95-246] using a morphological systematics approach. Handley employed 17 morphological characters and identified only four characters that reliably differentiated the three species he recognized, C. townsendii, C. rafinesquii, and C. mexicanus. All three species have been shown to lead relatively sedentary lives and have a wing morphology that limits long-distance dispersal. Further, populations of two species, C. townsendii and C. rafinesquii, are considered to be declining. Handley [Handley, C.O., 1959. A revision of the American bats of the genera Euderma and Plecotus. Proc. U.S. Nat. Mus. 110, 95-246] recognized five subspecies of C. townsendii (C. t. australis, C. t. ingens, C. t. pallescens, C. t. townsendii, and C. t. virginianus) and two of C. rafinesquii (C. r. rafinesquii and C. r. macrotis). Two C. townsendii subspecies, C. t. ingens and C. t. virginianus, are listed as endangered under the Endangered Species Act. These facts and the lack of a thorough molecular systematic examination of this genus were the impetus for this study. Using mitochondrial and nuclear intron DNA sequences, a molecular phylogeny was inferred. The combined DNA phylogeny supports Handley's [Handley, C.O., 1959. A revision of the American bats of the genera Euderma and Plecotus. Proc. U. S. Nat. Mus. 110, 95-246] designation of three species. Further the endangered subspecies, C. t. ingens and C. t. virginianus are corroborated, as were the monophyly of the other subspecies. However, the geographical ranges of two of these subspecies, C. t. pallescens and C. t. townsendii, are revised based on biogeographic distributions as understood from our results. Estimates of timing of divergences indicate that the three species, C. townsendii, C. rafinesquii, and C. mexicanus may have diverged before Pleistocene climatic oscillations began, and therefore their cladogenesis was not the result of those processes. However, subspecific divergences within C. townsendii appear to have occurred during and been driven largely by the climatic processes of the Pleistocene Epoch. We propose new hypotheses of dispersal scenarios that may have led to the current biogeography of these lineages.     

Redescription of Fabriciola tonerella Banse, 1959, and a new record of Novafabricia infratorquata (Fitzhugh, 1983) from the Mediterranean Sea, with a key for the Fabriciinae (Annelida: Polychaeta) of the Mediterranean Sea and the north-east Atlantic

The fabriciin sabellid species Fabriciola tonerella Banse, 1959 living on gastropod shells inhabited by hermit crabs collected in the western Mediterranean Sea is redescribed. Another species of Fabriciinae, Novafabricia infratorquata (Fitzhugh, 1983), is recorded for the first time from the Mediterranean Sea and found in the same habitat is described as well. The variability of characters depending on size and geographical distribution is given for both species. Moreover, selected characters for all known species of Fabriciola and Novafabricia as well as a simple key for all Fabriciinae of the Mediterranean Sea and the north-east Atlantic are provided.     

The number of segregating genes implied by continuous variation

Phenotypic distributions indistinguishable from normal distributions are often interpreted as showing the segregation of many genes. Such characters may then be passed over by physiologists and developmental biologists as too complex or otherwise unsuitable for developmental studies. In this paper we present several simple genetic models in which only 2 or 3 loci account for the great majority of the variance, and we show that even with large sample sizes, such distributions may be indistinguishable from normal distributions. The conclusion that a normal distribution must reflect the segregation of a large number of loci is therefore not necessarily justified. Hopefully, problems of the development of quantitative characters which were previously thought to be unapproachable may now be seen to be open to critical examination.This work was supported by a grant from the Science Research Council.     

Analysis of the diet of Natterer's bat Myotis nattereri and the common long-eared bat Plecotus auritus in the West of Ireland

The diets of Natterer's bat Myotis nattereri and the common long-eared bat Plecotus auritus were investigated at a nursery colony of each species by analysis of droppings collected monthly from May to September. Respectively, 68% and 42% of the diets comprised items presumed to have been gleaned from foliage or other surfaces: diurnal insects, insects which rarely fly, and non-flying arthropods. Such surfaces included the ground: centipedes were eaten by both bats, and the muscid, Scatophaga stercoraria , usually associated with cattle dung, was a common prey. Indeed, the prevalence of cattle-farming around both of the bat roosts almost certainly contributed indirectly, in the form of S. stercoraria , to a significant part of the Diptera consumed. The chief food of M. nattereri was the larger Diptera, but Trichoptera, Hymenoptera and Arachnida were also important. Lepidoptera, Coleoptera and Hemiptera were minor prey, with Dermaptera and Chilopoda taken occasionally. Diptera, closely followed by Lepidoptera, together accounted for nearly two-thirds of the diet of P. auritus. Also taken, in descending order of importance, were Trichoptera, Arachnida, Chilopoda, Coleoptera, Dermaptera, Hymenoptera and Hemiptera. Plecotus auritus , but not M. nattereri , was evidently able to take a few small insects such as aphids and lesser nematoceran Diptera.