Name: ARTURO BENINCA MARTINELLI
Type: PhD thesis
Publication date: 22/02/2018
Advisor:

Name	Role
Celso Oliveira Azevedo	Co-advisor *
Valéria Fagundes	Advisor *

Examining board:

Name	Role
Claudio José Barros de Carvalho	External Examiner *
Gustavo Rocha Leite	Internal Examiner *
ISABEL DE CONTE CARVALHO DE ALENCAR	External Alternate *
Kátia Cristina Machado Pellegrino	External Examiner *
Leonora Pires Costa	Internal Alternate *
Valéria Fagundes	Advisor *
Yuri Luiz Reis Leite	Internal Examiner *

Summary: The study of geographical distribution of living beings is a simple and accurate definition for
biogeography. In this discipline, descriptive and interpretive perspectives are found, in which the
first describes taxa distribution patterns and the second, explanations for found patterns. The
current thesis was built in a context of working with a cosmopolitan group WHERE patterns and
biogeographical processes which made the respective distribution are not known. This is the
case of Dissomphalus Ashmead, 1893 (Hymenoptera Bethylidae), a gender of parasitoid
wasps, found in all zoogeographical regions. Everything points that the group is old, once the
molecular dating and the Behylidae fossil presence suggest the antiquity of the family matching
the Gondwana fragmentation. In this case, it was applied to Dissomphalus the “Gondwana
Paradigm” as possible explanation for cosmopolitan distribution. Hypotheses were considered
that Dissomphalus phylogenetic history and age are related to Gondwana fragmentation
sequence. The thesis was arranged in four chapters which consist the steps for biogeographical
knowledge construction of the group, that goes since the setting of geographical distribution
patterns to hypothesis’ effective tests. The first step, determination of distribution patterns, was
the subject of the first two projects and identified by Panbiogeography means. Chapter 1
approaches potential solutions for quantitative congruency, which currently are methodological
setbacks for panbiogeography. The distribution points of Dissomphalus were used as models, in
which were georeferenced in 10.191 testimonies of published group so far to November/16 and
chosen 7.457 concerning American specimens. The Parsimonious Analysis of Endemicity with
Progressive Characters Elimination (PAE-PCE) was adopted for general trace selection, in
which the method was improved by the analysis of multiple spatial scale. 20 grates of grid sizes
were used, varying from 0.5º x 0.5º (1.0º to 10º) and another grate which geographical units of
analysis the biogeographical provinces of Neotropic were. All scales were analyzed and
compared. The application of multiple scales allowed the resolution of most part of the
congruency problems in quantitative panbiogeography pointed to identification of generalized
trace. In Chapted 2, PAE-PCE improved method was applied to effectively identify distribution
patterns of American Dissomphalus. Six areas of general trac were identified (Amazon, Atlantic
forest, Mesoamerica, Northern Andes, Lesser Antilles and Yungas) and three major
panbiogeographical nodes, here referred as supertracks. All generalized tracks were correlated
to historical events of Americas, which revealed to be related to tectonism occurred from
Miocene (~23 Million years), period in which coincides to neotectonic scope. In this case, in
Chapter 3, a neotectonic model was proposed with potential to explain distribution patterns of
Dissomphalus species, and, also, of another biological groups, specially the ones in Atlantic
forest. The built model was based in the possibility of weakening crust zones being reactivated
along geological time, thus offering embossed modifications capable of changing the
populations’ genic flow of many specimens and generating the found patterns. In Chapter 4,
biogeographic hypothesis for Dissomphalus were tested by phylogenic and molecular dating.
178 specimens of Dissomphalus were selected of 30 countries which englobes all
zoogeographic regions, except the Sino-Japanese. Five molecular markers were sequenced,
being one mitochondrial (COI) and four nuclear (28S, LW, POL2 and EFa2), inferred molecular
dating, phylogenetic trees and ancient area estimatives. Four major lineages of Dissomphalus
were identified, of which was not possible to obtain with support the relation between them:
Brand New-New World, Afromadagascan, Afroasian and Oriental. In general, the paradigm of
Gondwana was refuted to Dissomphalus and the panbiogeographical correlations of Chapter 2,
corroborated. The group revealed to be related to Eocene-Paleocene (46,65 Million years
[33,55-62,79]) with diversification in Miocene, which are more recent period than Gondwana’s
fragmentation. Seemingly, the group colonized the world from east to west, beginning in Africa.
Biogeographical dispersion processes were corroborated by molecular dating to explain the
cosmopolitanism of the group. Dissomphalus disperse to west of Africa through the oceans by
rafts in two distinct moments, first reaching the New World (Oligocene-Eocene) and more
recently the Brand-New World (Miocene). The group also went through terrestrial bridges that
facilitated the dispersion, as such the Panama Isthmus (Miocene and Pliocene) and Horn Of
Africa-Arabian Peninsula (Miocene and Pliocene), both, as it appears, at least twice in distinct
period.
Keywords: Biogeography, Dispersal, Vicariance, South America, Atlantic Forest, Neotectonics

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