Christophe Daugeron

Phylogenetics and Ecology: As Many Characters as Possible Should Be Included in the Cladistic Analysis☆

As many data as possible must be included in any scientific analysis, provided that they follow the logical principles on which this analysis is based. Phylogenetic analysis is based on the basic principle of evolution, i.e., descent with modification. Consequently, ecological characters or any other nontraditional characters must be included in phylogenetic analyses, provided that they can plausibly be postulated heritable. The claim of Zrzavý (1997, Oikos 80, 186–192) or Luckow and Bruneau (1997, Cladistics 13, 145–151) that any character of interest should be included in the analysis is thus inaccurate. Many characters, broadly defined or extrinsic (such as distribution areas), cannot be considered as actually heritable. It is argued that we should better care for the precise definition and properties of characters of interest than decide a priori to include them in any case in the analysis. The symmetrical claim of de Queiroz (1996, Am. Nat. 148, 700–708) that some characters of interest should better be excluded from analyses to reconstruct their history is similarly inaccurate. If they match the logical principles of phylogenetic analysis, there is no acceptable reason to exclude them. The different statistical testing strategies of Zrzavý (1997) and de Queiroz (1996) aimed at justifying inclusion versus exclusion of characters are ill‐conceived, leading respectively to Type II and Type I errors. It is argued that phylogenetic analyses should not be constrained by testing strategies that are downstream of the logical principles of phylogenetics. Excluding characters and mapping them on an independent phylogeny produces a particular and suboptimal kind of secondary homology, the use of which can be justified only for preliminary studies dealing with broadly defined characters.

Are empidine dance flies major flower visitors in alpine environments? A case study in the Alps, France.

Pollination is one of the most important ecosystem services and bees the most important pollinators. As a population decline of bees has been documented in numerous regions of the world, it is crucial to develop understanding on other possible pollinators. Here, we study the potential pollination impact of Diptera, and among them Empidinae, in an alpine environment, where the abundance of bees is naturally lower. Interactions between 19 entomophilous plants and their flower visitors were recorded in a subalpine meadow in the French Alps during six weeks. Visitation frequencies were used to build the flower–visitor network. Our results show that interactions between flies and plants are dominant; flies represent more than 60% of all visitors, with 54% of them being Empidinae. We especially found that flies, Empidinae and bees are the main visitors of 11, three and one plants, respectively. When considering both bees and Syrphidae together, six plants were more visited by Empidinae; when considering bees and Syrphidae separately, 10 plants were more visited by Empidinae than by bees or Syrphidae. The results support the idea that flies widely replace bees as main flower visitors at altitude, and among them the Empidinae might play a key role in pollination.

Extreme male leg polymorphic asymmetry in a new empidine dance fly (Diptera: Empididae).

A new dance fly (Empididae: Empidinae) with hugely modified male fore tarsus, either on the right, left, both or neither sides, is described from Japan. Such massive polymorphic asymmetry occurring with so high an incidence in a population is previously unreported. In view of the courtship behaviour of other Empidinae, we hypothesize that the oversized tarsus is a secondary sexual character employed by males for attracting females. Alternative hypotheses are also discussed. We suggest that this extraordinary new species is a potential model for the study of mating biology in Empidinae and the evolution of mating systems in general.

Organization, usefulness and limitations of an ATBI (All Taxa Biodiversity Inventory): the inventory of terrestrial invertebrates in the Mercantour National Park

ABSTRACT We present here the objectives, organization and preliminary results of the invertebrate inventory of the Mercantour National Park (“Terrestrial Invertebrates Module” of the ATBI Mercantour/Alpi Marittime), carried out by 68 scientists and field-collectors from 2009 to 2012. The conceptual framework of an ATBI and the interactions between inventorying and frequently associated research aims are discussed. The sampling strategy adopted combined standardized multi-taxon sampling (continuous and occasional) with individual taxon-centred sampling. The successive tasks of field sampling, specimen coding, sorting and identification, molecular analysis and data management are briefly described. A coherent system of sample coding ensured the traceability of specimens. The project results in terms of sampling effort, abundance of collected invertebrate fauna and biological diversity are presented globally, by techniques, by sites and by higher taxonomic groups: 259,412 specimens were obtained from 761 sorted samples (out of 1243 samples collected), belonging to 1725 identified species and subspecies, in addition to which 37 species new to science have been recognized. These data are progressively being integrated into the CardObs database of the INPN and are partly accessible through the EDIT and GBIF websites. Molecular analyses (barcoding) have provided 2800 sequences (mostly for the COI gene in Gastropoda, Collembola and Hymenoptera). Analysis of these sequences indicates a significant amount of cryptic diversity in the material. From a macro-ecological perspective, endemicity in the park decreases globally from south to north and species richness decreases at higher elevations, but the data are uneven and patterns vary according to group. Based on this four-year experience, it appears that a smaller survey area with lower habitat diversity would be better suited to the aims of an ATBI when the available resources are limited, and would enhance the efficiency of collective work in the field.

A new species of Phyllodromia Zetterstedt, 1837 (Insecta, Diptera, Empididae, Hemerodromiinae) from Vanuatu

Plant A. R. & Daugeron C. 2009. — A new species of Phyllodromia Zetterstedt, 1837 (Insecta, Diptera, Empididae, Hemerodromiinae) from Vanuatu. Zoosystema 31 (3): 519-524. ABSTRACT Phyllodromia variabilis n. sp. (Diptera, Empididae, Hemerodromiinae) is described from Espiritu Santo, Vanuatu and provisionally assigned to Phyllodromia Zetterstedt, 1837. Systematic relationships with other Phyllodromia species and with the closely related genus Chelipoda Macquart, 1823 are discussed and it is concluded that the new species is probably more closely related to species of Phyllodromia from New Zealand than those of the Northern Hemisphere.

Phylogenetic systematics of the Gondwanan Empis macrorrhyncha group (Diptera, Empididae, Empidinae)

The Empis macrorrhyncha group (Diptera: Empididae) from cool to warm temperate areas of South America and Australia is diagnosed and cladistically analysed, and five new species, Empis animosasp.n., E. austerasp.n., E. maculosasp.n., E. occidentalissp.n. and E. pedivillosulasp.n., are described. Cladistic analysis of 23 adult morphological characters for 14 species of the group generated a single tree of 28 steps (CI = 0.82; RI = 0.93). Monophyly was established on the basis of a single apomorphy, possession of a bilobed cercus of the male hypopygium. Three main clades were inferred: clade 1 included three Patagonian and a single southwestern Australian species; clade 2 included two species from southeastern Australia; clade 3 included a large Patagonian group of five species and a single southeastern Australian species. The E. fulvicollis complex (clade 1) is a sister‐group of the E. macrorrhyncha complex (clades 2 + 3). A provisional historical biogeographic hypothesis is advanced correlating the appearance of the South American and Australian sister lineages with the timing of the break‐up of Gondwana.

The Mercantour/Alpi Marittime All Taxa Biodiversity Inventory (ATBI): achievements and prospects

ABSTRACT This paper summarizes the main results from taxonomic inventories drawn up as part of the ATBI of the Mercantour/Alpi Marittime parks in the French and Italian Alps. It compares the data on different groups of the invertebrate fauna published in the Zoosystema issues devoted to the ATBI with those recorded in the EDIT (European Distributed Institute of Taxonomy) and INPN (Inventaire national du Patrimoine naturel) ATBI databases. It also highlights the contribution of the ATBI inventory to a better knowledge of the biology and ecology of various animal taxa, as well as to the use of the data in conservation management. Overall, 927 animal taxa and 38 host plants are documented in the two special issues, of which 904 are recorded from both parks and 400 are new for one or both parks. Twelve species (ten terrestrial and two aquatic) and one genus (aquatic) are described as new for science, this increases to 30 the number of new invertebrate taxa described since 2006 from the Mercantour/Alpi Marittime parks. However, due to taxonomic uncertainty about material sorted to morphospecies and delays in species descriptions, these represent only a fraction of the taxa recognized as new during the ATBI. This faunal inventory, which includes 88% insect taxa, greatly enhances knowledge of Mercantour/Alpi Marittime biodiversity, notably for several poorly studied taxonomic groups. It also confirms the importance of the area as a European biodiversity hotspot, especially for arthropods and the groundwater fauna, by highlighting the remarkable percentage of endemic species, some of which are rare and poorly documented in the literature.

Altitudinal, temporal and trophic partitioning of flower-visitors in Alpine communities

The cross-pollination of most alpine plants depends on insects, whose altitudinal distribution is limited by temperature. However, although global warming is causing shifts in temporal and spatial species distribution, we are still largely unaware of how plant-pollinator interactions change with elevation and time along altitudinal gradients. This makes the detection of endangered interactions and species challenging. In this study, we aimed at providing such a reference, and tested if and how the major flower-visiting insect orders and families segregated by altitude, phenology and foraging preferences along an elevational gradient from 970 m to 2700 m in the Alps. Flies were the main potential pollinators from 1500 m, as bees and beetles decreased rapidly above that limit. Diptera, Coleoptera and Hymenoptera differed significantly in the angiosperm assemblages visited. Within Diptera, the predominant group, major families segregated by both phenology and foraging preferences along the gradient. Empidids, muscids and anthomyiids, whose role in pollination has never been investigated, dominated the upper part of the gradient. Our results thus suggest that flies and the peculiar plants they visit might be particularly at risk under global warming, and highlight the blatant lack of studies about critical components of these rich, yet fragile mountain ecosystems.

Phylogenetic reappraisal and taxonomic review of the Empis (Coptophlebia) hyalipennis-group (Diptera : Empididae : Empidinae)

The Empis (Coptophlebia) hyalipennis-group is redefined, keyed, cladistically analysed and three new species, Empis doi, sp. nov., Empis pseudohystrichopyga, sp. nov. and Empis pseudovillosipes, sp. nov., are described (from Thailand, China and Taiwan respectively). A cladistic analysis of 27 adult morphological characters was performed for Empis hyalipennis (the type-species of the subgenus Coptophlebia), and 13 Nearctic, Oriental and Palaearctic species hypothesised as being closely related, which resulted in a single tree of 31 steps (CI = 0.96; RI = 0.98). Monophyly of the group was established on the basis of two apomorphic characters of the male hypopygium. Based on these results and previous studies of the Empis-Coptophlebia complex of subgenera within the genus Empis, the E. (C.) hyalipennis-group appears to be much richer in species, exhibit greater morphological heterogeneity and be more widely distributed than expected. It is strongly recommended that any future revision of the subfamily Empidinae should apply the name Coptophlebia only to the monophyletic Empis (Coptophlebia) hyalipennis-group defined in this study.

Where do the Neotropical Empidini lineages (Diptera: Empididae: Empidinae) fit in a worldwide context?

The tribe Empidini (Diptera: Empididae: Empidinae) is a diverse group with fourteen genera, seven of which are exclusive to the Neotropical region: Bolrhamphomyia Rafael, Chilerhamphomyia Rafael, Hystrichonotus Collin, Lamprempis Wheeler and Melander, Macrostomus Wiedemann, Opeatocerata Melander and Porphyrochroa Melander. Although Empidini itself is likely paraphyletic, many presumably monophyletic genera and species groups are recognized. Here, we apply DNA sequences from multiple genes to infer the phylogeny of Empidini, focusing on placing the Neotropical lineages within the entire tribe and identifying monophyletic groups. We included 98 Empidini taxa along with 18 outgrous terminals, spanning the diversity within the group. The results from the analyses performed are largely similar, with major groupings of genera in common. Specifically, the analyses recovered a monophyletic Hilarini and a paraphyletic Empidini. Most species from Chile and Argentina (Andean region) are found to belong to an early branching lineage within Empidinae, and are not monophyletic with other Empidini. A large portion of the remaining Neotropical Empidini (not Andean) comprises a single clade that includes four endemic genera and a number of Neotropical Empis Linneaus species. Macrostomus and Porphyrochroa each recovered as monophyletic and sister to one another, although generic placement of a few taxa remains uncertain due to conflicting morphological features. Lamprempis+Opeatocerata are also found to be sister-taxa in most analyses. Several large genera were found to be polyphyletic or paraphyletic including Empis and Rhamphomyia Meigen. We evaluate our findings and discuss them in light of current Empidinae taxonomy.