Francisco Hita Garcia

Forest Fragmentation and Selective Logging Have Inconsistent Effects on Multiple Animal-Mediated Ecosystem Processes in a Tropical Forest

Forest fragmentation and selective logging are two main drivers of global environmental change and modify biodiversity and environmental conditions in many tropical forests. The consequences of these changes for the functioning of tropical forest ecosystems have rarely been explored in a comprehensive approach. In a Kenyan rainforest, we studied six animal-mediated ecosystem processes and recorded species richness and community composition of all animal taxa involved in these processes. We used linear models and a formal meta-analysis to test whether forest fragmentation and selective logging affected ecosystem processes and biodiversity and used structural equation models to disentangle direct from biodiversity-related indirect effects of human disturbance on multiple ecosystem processes. Fragmentation increased decomposition and reduced antbird predation, while selective logging consistently increased pollination, seed dispersal and army-ant raiding. Fragmentation modified species richness or community composition of five taxa, whereas selective logging did not affect any component of biodiversity. Changes in the abundance of functionally important species were related to lower predation by antbirds and higher decomposition rates in small forest fragments. The positive effects of selective logging on bee pollination, bird seed dispersal and army-ant raiding were direct, i.e. not related to changes in biodiversity, and were probably due to behavioural changes of these highly mobile animal taxa. We conclude that animal-mediated ecosystem processes respond in distinct ways to different types of human disturbance in Kakamega Forest. Our findings suggest that forest fragmentation affects ecosystem processes indirectly by changes in biodiversity, whereas selective logging influences processes directly by modifying local environmental conditions and resource distributions. The positive to neutral effects of selective logging on ecosystem processes show that the functionality of tropical forests can be maintained in moderately disturbed forest fragments. Conservation concepts for tropical forests should thus include not only remaining pristine forests but also functionally viable forest remnants.

A Preliminary Checklist of the Ants (Hymenoptera: Formicidae) of Kakamega Forest (Kenya)

ABSTRACT A preliminary species checklist of the ants (Hymenoptera: Formicidae) of Kakamega Forest, Western Kenya, is presented. The species list is based on specimens sampled from 1999 until 2009, which are deposited in the ant collection of the Zoological Research Museum Koenig, Bonn, Germany, and the Natural History Museum of Los Angeles County, Los Angeles, United States. The known ant fauna comprises 11 subfamilies with 52 genera and 288 species. This species richness is the second highest reported from the Afrotropical zoogeographical region. The observed ant fauna can be divided into a group of forest habitat species with approximately 180 species and another group, which consists of about 100 species, of open habitat specialists. The greatest part of the ant fauna (204 species or 71%) could be identified to species level but the rest remains unidentifiable or undescribed (84 species or 29%). The preliminary data from the observed ant fauna seems to support the hypothesis that Kakamega Forest is the eastern-most remnant of the former Guineo-Congolian rain forest belt, while the zoogeographical influence from East African Afromontane forests seems to be less significant.

Improved Phylogenetic Analyses Corroborate a Plausible Position of Martialis heureka in the Ant Tree of Life

Martialinae are pale, eyeless and probably hypogaeic predatory ants. Morphological character sets suggest a close relationship to the ant subfamily Leptanillinae. Recent analyses based on molecular sequence data suggest that Martialinae are the sister group to all extant ants. However, by comparing molecular studies and different reconstruction methods, the position of Martialinae remains ambiguous. While this sister group relationship was well supported by Bayesian partitioned analyses, Maximum Likelihood approaches could not unequivocally resolve the position of Martialinae. By re-analysing a previous published molecular data set, we show that the Maximum Likelihood approach is highly appropriate to resolve deep ant relationships, especially between Leptanillinae, Martialinae and the remaining ant subfamilies. Based on improved alignments, alignment masking, and tree reconstructions with a sufficient number of bootstrap replicates, our results strongly reject a placement of Martialinae at the first split within the ant tree of life. Instead, we suggest that Leptanillinae are a sister group to all other extant ant subfamilies, whereas Martialinae branch off as a second lineage. This assumption is backed by approximately unbiased (AU) tests, additional Bayesian analyses and split networks. Our results demonstrate clear effects of improved alignment approaches, alignment masking and data partitioning. We hope that our study illustrates the importance of thorough, comprehensible phylogenetic analyses using the example of ant relationships.

The hyper-diverse ant genus Tetramorium Mayr (Hymenoptera, Formicidae) in the Malagasy region ‑ taxonomic revision of the T. naganum, T. plesiarum, T. schaufussii, and T. severini species groups

Abstract The taxonomy of the Tetramorium naganum, T. plesiarum, T. schaufussii, and T. severini species groups are revised for the Malagasy region. A total of 31 species are treated, of which 22 are newly described and nine redescribed. This increases the richness of the hyper-diverse genus Tetramorium in the Malagasy region to 106 species, which makes it the most species-rich genus in the region. Twenty-nine of the treated species are endemic to Madagascar, one is endemic to the Comoros, and one species is found predominantly in Madagascar but also on the island of Reunion. The T. naganum species group contains five species, which are mainly distributed in the rainforests and montane rainforests of eastern and northern Madagascar: T. alperti sp. n., T. dalek sp. n., T. enkidu sp. n., T. gilgamesh sp. n., and T. naganum Bolton, 1979. The T. plesiarum species group holds five species: T. bressleri sp. n., T. hobbit sp. n., T. gollum sp. n., T. mars sp. n., and T. plesiarum Bolton, 1979. All five are arid-adapted species occurring in the southwest and west of Madagascar. The second-most species-rich group in the region is the T. schaufussii species group with 20 species, most of which inhabit rainforests or montane rainforests of eastern and northern Madagascar. This group includes two species complexes each containing ten species: the T. cognatum complex with the species T. aspis sp. n., T. camelliae sp. n., T. cognatum Bolton, 1979, T. freya sp. n., T. gladius sp. n., T. karthala sp. n., T. myrmidon sp. n., T. proximum Bolton, 1979, T. rumo sp. n., and T. tenuinode sp. n.; and the T. schaufussii complex with the species T. merina sp. n., T. monticola sp. n., T. nassonowii Forel, 1892 stat. n., T. obiwan sp. n., T. pseudogladius sp. n., T. rala sp. n., T. schaufussii Forel, 1891, T. sikorae Forel, 1892 (= T. latior (Santschi, 1926)), T. scutum sp. n., T. xanthogaster Santschi, 1911. The last group treated in this study is the T. severini species group, which contains only the species T. severini (Emery, 1895). This very conspicuous species is widely distributed in the rainforests and montane rainforests of eastern and northern Madagascar. All four groups are fully revised with group diagnoses, illustrated species-level identification keys, and detailed descriptions for all species that include multifocused montage images and distribution maps.

The Ants of Kenya (Hymenoptera: Formicidae)—Faunal Overview, First Species Checklist, Bibliography, Accounts for All Genera, and Discussion on Taxonomy and Zoogeography

ABSTRACT This study represents the first comprehensive treatment of Kenyan myrmecofauna, which includes a faunal overview, the first species checklist for Kenya, a bibliography, illustrated accounts for all genera, and discussions on taxonomy and biogeography. The checklist is based on material collected from several important sampling localities and literature records. The main localities include Kakamega Forest, a tropical rainforest in Western Kenya, Mpala Research Centre, a savannah region in Laikipia, Central Kenya, and Arabuko Sokoke Forest, a coastal tropical dry forest. Currently, the country is known to hold 596 species and 63 genera from 12 subfamilies. The most diverse subfamilies are Myrmicinae (311 species, 26 genera), Formicinae (107 species, 12 genera), and Ponerinae (75 species, 10 genera). Additional collections from further localities will likely increase these counts and probably reach more than 650 species. Based on the examined material, the Kenyan ant fauna consists of distinct regional elements with different Zoogeographic affinities and minimal overlap.

New records of ant species from Yunnan, China

Abstract As with many other regions of the world, significant collecting, curation, and taxonomic efforts will be needed to complete the inventory of China’s ant fauna. This is especially true for the highly diverse tropical regions in the south of the country, where moist tropical forests harbor high species richness typical of the Southeast Asian region. We inventoried ants in the Xingshuangbanna prefecture, Yunnan, in June 2013, using a variety of methods including Winkler extraction and hand collection to sample ant diversity. We identified 213 species/morphospecies of ants from 10 subfamilies and 61 genera. After identification of 148 valid species of the 213 total species collected, 40 species represent new records for Yunnan province and 17 species are newly recorded for China. This increases the total number of named ant species in Yunnan and China to 447 and 951 respectively. The most common species collected were Brachyponera luteipes and Vollenhovia emeryi. Only one confirmed exotic species Strumigenys membranifera, was collected, although several others were potentially introduced by humans. These results highlight the high biodiversity value of the region, but also underscore how much work remains to fully document the native myrmecofauna.

X-Ray microtomography for ant taxonomy: An exploration and case study with two new Terataner (Hymenoptera, Formicidae, Myrmicinae) species from Madagascar

We explore the potential of x-ray micro computed tomography (μCT) for the field of ant taxonomy by using it to enhance the descriptions of two remarkable new species of the ant genus Terataner: T. balrog sp. n. and T. nymeria sp. n.. We provide an illustrated worker-based species identification key for all species found on Madagascar, as well as detailed taxonomic descriptions, which include diagnoses, discussions, measurements, natural history data, high-quality montage images and distribution maps for both new species. In addition to conventional morphological examination, we have used virtual reconstructions based on volumetric μCT scanning data for the species descriptions. We also include 3D PDFs, still images of virtual reconstructions, and 3D rotation videos for both holotype workers and one paratype queen. The complete μCT datasets have been made available online (Dryad, https://datadryad.org) and represent the first cybertypes in ants (and insects). We discuss the potential of μCT scanning and critically assess the usefulness of cybertypes for ant taxonomy.

Taxonomy of the ant genus Proceratium Roger (Hymenoptera, Formicidae) in the Afrotropical region with a revision of the P.arnoldi clade and description of four new species.

Abstract The taxonomy of the genus Proceratium Roger is updated for the Afrotropical region. We give an overview of the genus in the region, provide an illustrated identification key to the three clades (Proceratium arnoldi, Proceratium stictum and Proceratium toschii clades) and revise the Proceratium arnoldi clade. Four new species from the Proceratium arnoldi clade are described as new: Proceratium sokoke sp. n. from Kenya, Proceratium carri sp. n. from Mozambique, and Proceratium nilo sp. n. and Proceratium sali sp. n. from Tanzania. In order to integrate the new species into the existing taxonomic system we present an illustrated identification key to distinguish the seven Afrotropical species of the Proceratium arnoldi clade. In addition, we provide accounts for all members of the Proceratium arnoldi clade including detailed descriptions, diagnoses, taxonomic discussions, distribution data and high quality montage images.

The Tetramorium tortuosum species group (Hymenoptera, Formicidae, Myrmicinae) revisited - taxonomic revision of the Afrotropical T. capillosum species complex

Abstract In this study we revise the taxonomy of the Tetramorium tortuosum species group members encountered in the Afrotropical region, which we have placed in its own subgroup: the Tetramorium capillosum species complex. We re-describe the two previously known species Tetramorium capillosum Bolton and Tetramorium tabarum Bolton, and describe the new species Tetramorium hecate sp. n. The geographic distribution of the three species appears to be restricted to the equatorial rainforests of Central Africa. We provide a diagnosis of the Tetramorium capillosum species complex, an illustrated identification key to species level, and worker-based species descriptions, which include diagnoses, discussions, high-quality montage images, and distribution maps. Furthermore, we discuss biogeography and composition of the globally distributed Tetramorium tortuosum group.

Next-generation morphological character discovery and evaluation: an X-ray micro-CT enhanced revision of the ant genus Zasphinctus Wheeler (Hymenoptera, Formicidae, Dorylinae) in the Afrotropics.

Abstract New technologies for imaging and analysis of morphological characters offer opportunities to enhance revisionary taxonomy and better integrate it with the rest of biology. In this study, we revise the Afrotropical fauna of the ant genus Zasphinctus Wheeler, and use high-resolution X-ray microtomography (micro-CT) to analyse a number of morphological characters of taxonomic and biological interest. We recognise and describe three new species: Z. obamai sp. n., Z. sarowiwai sp. n., and Z. wilsoni sp. n. The species delimitations are based on the morphological examination of all physical specimens in combination with 3D scans and volume reconstructions. Based on this approach, we present a new taxonomic discrimination system for the regional fauna that consists of a combination of easily observable morphological characters visible at magnifications of around 80–100 ×, less observable characters that require higher magnifications, as well as characters made visible through virtual dissections that would otherwise require destructive treatment. Zasphinctus are rarely collected ants and the material available to us is comparatively scarce. Consequently, we explore the use of micro-CT as a non-invasive tool for the virtual examination, manipulation, and dissection of such rare material. Furthermore, we delineate the treated species by providing a diagnostic character matrix illustrated by numerous images and supplement that with additional evidence in the form of stacked montage images, 3D PDFs and 3D rotation videos of scans of major body parts and full body (in total we provide 16 stacked montage photographs, 116 images of 3D reconstructions, 15 3D rotation videos, and 13 3D PDFs). In addition to the comparative morphology analyses used for species delimitations, we also apply micro-CT data to examine certain traits, such as mouthparts, cuticle thickness, and thoracic and abdominal muscles in order to assess their taxonomic usefulness or gain insights into the natural history of the genus. The complete datasets comprising the raw micro-CT data, 3D PDFs, 3D rotation videos, still images of 3D models, and coloured montage photos have been made available online as cybertypes (Dryad, http://dx.doi.org/10.5061/dryad.4s3v1).